ArticlePDF Available

Two new subterranean-adapted pseudoscorpions (Pseudoscorpiones: Neobisiidae: Parobisium) from Beijing, China

Authors:

Abstract and Figures

Two new troglomorphic pseudoscorpion species, Parobisium magangensis sp. n. and P. yuantongi sp. n., belonging to the family Neobisiidae, are described based on specimens collected in karst caves from Beijing, China. These are the first troglomorphic pseudoscorpions discovered from caves in northern China. Detailed diagnosis, descriptions, and illustrations are provided. We also offer future research and management recommendations for these two new pseudoscorpion species.
Content may be subject to copyright.
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by D. Harms: 2 Aug. 2019; published: 27 Aug. 2019 145
Zootaxa 4661 (1): 145–160
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4661.1.7
http://zoobank.org/urn:lsid:zoobank.org:pub:1239FB1C-0E57-4493-8236-079ADAD99918
Two new subterranean-adapted pseudoscorpions
(Pseudoscorpiones: Neobisiidae: Parobisium) from Beijing, China
ZEGANG FENG1, J. JUDSON WYNNE2 & FENG ZHANG1,3
1The Key Laboratory of Invertebrate Systematics and Application, College of Life Sciences, Hebei University, Baoding, Hebei 071002,
P. R. China
2Department of Biological Sciences, Merriam-Powell Center for Environmental Research, Northern Arizona University, Flagstaff,
Arizona 86011, U.S.A.
3Corresponding author. E-mail: dudu06042001@163.com
Abstract
Two new troglomorphic pseudoscorpion species, Parobisium magangensis sp. n. and P. yuantongi sp. n., belonging
to the family Neobisiidae, are described based on specimens collected in karst caves from Beijing, China. These are
the first troglomorphic pseudoscorpions discovered from caves in northern China. Detailed diagnosis, descriptions,
and illustrations are provided. We also offer future research and management recommendations for these two new
pseudoscorpion species.
Key words: Parobisium, troglobionts, cavernicolous, taxonomy, false scorpions
Introduction
The distribution of karst (a dissolution substrate including limestone) in China is extensive and there are many
karstic regions with a poorly known fauna (Zhang & Zhu 2012). Most studies of cave-dwelling arthropods have fo-
cused on the extensive South China Karst, which encompasses four administrative provinces in southern China and
contains thousands of caves. To date, at least 301 cave-dwelling arthropod species have been identified from this
region (Tian & Clarke 2012; Ran & Yang 2015; Gao et al. 2018; Liu & Wynne 2019). Our knowledge of northern
cave-dwelling arthropods is limited primarily to the Beijing area. To date, Tong & Li (2008) reported six species of
cave-dwelling spiders from Beijing and adjacent areas.
Until now, there have not been any documented cases of subterranean-adapted pseudoscorpions in northern
China, and our knowledge has been limited to 20 troglomorphic pseudoscorpion species (Families Chthoniidae,
Neobisiidae and Chernetidae) from southern China (Schawaller 1995; Mahnert 2003, 2009; Mahnert & Li 2016;
Gao et al. 2017; Li et al. 2017; Gao et al. 2018).
In his global catalogue, Harvey (2013) reported 16 species of Parobisium but three troglobitic species from Chi-
na (P. martii Mahnert, 2003, P. scaurum Mahnert, 2003 and P. titanium Mahnert, 2003) were recently transferred to
Bisetocreagris Ćurčić, 1983 by Mahnert & Li (2016) while Guo & Zhang (2016) added two additional epigean spe-
cies (P. wangae Guo & Zhang, 2016 and P. xiaowutaicum Guo & Zhang, 2016) from China to Parobisium. Hence,
there are currently 15 species of Parobisium from China (two species), North America (seven species), Japan (four
species) and Korea (two species) (Harvey 2013; Guo & Zhang 2016). Of these, only three species are troglobitic:
P. yosemite Cokendolpher & Krejca, 2010 (Indian Cave, Yosemite National Park, California, U.S.A), P. charlotteae
Chamberlin, 1962 (Redmond Lava Cave, Deschutes County, Oregon, U.S.A), and P. anagamidense (Morikawa,
1957) (Anagami-d Cave, Kochi Prefecture, Japan).
In this study, we describe the first subterranean-adapted species of Parobisium from Northern China. All speci-
mens are troglomorphic and display typical adaptation to subterranean habitat such as larger body size, lack of eyes,
and long, slender pedipalps and legs. We also provide recommendations for future research and management to
protect these new pseudoscorpion species and their habitats.
FENG ET AL.
146 · Zootaxa 4661 (1) © 2019 Magnolia Press
Material and methods
Study area. Fangshan World Geopark is located southwest of Beijing, China and comprises an area of 953.95 km2
(Fig. 1). Featuring a peak cluster landscape (Lu 2007; Cao et al. 2014), this geopark is characterized by temperate
karst (Yu et al. 2011) and is composed principally of the Ordovician Majiagou Formation limestone and Mesopro-
terozoic Jixian Siluoshan siliceous strip dolomite (Lu et al. 2010). Due to neotectonic movement and climatic condi-
tions, the Fangshan landform and multi-layered karst caves are unique to the region (Lu et al. 2010). This geopark
supports at least 100 documented caves. Our study caves were selected based on information provided by local
villagers and the presence of sufficient depth to support deep zone conditions (defined below). Because these caves
have not been explored by any of the speleological groups working in China, cave maps were unavailable.
Magang Cave (Figs 1, 9) is located in a remote mountain ~2 km south of the town of Xiayunling. The surround-
ing area is largely forested and the closest agricultural fields/ human settlements are approximately 1 km away. This
limestone cave has one triangular entrance (~2 meters high and 0.2 to 0.4 meters wide) and is approximately 200
meters in length, and extends horizontally. The last ~120 m of this cave support active speleothems.
Shenxian Cave (Figs 1, 10) is located 1.2 km southwest of the town of Jinjitai. The cave is situated near a pig
farm and within low scrub vegetation. This cave is approximately 100 meters in length, has a dome shaped entrance
(approximately two meters wide), and extends horizontally.
Field sampling. Cave-dwelling arthropods were collected from the two caves on June 10, 2018. Observers
conducted a direct intuitive search (Wynne et al. 2019) of arthropods from the entrance area to the deep zone of
these caves by scanning the cave floor, ceiling, and walls. For the floor, observers examined the space between the
gravel and the ground, and searched beneath rocks (Figs 9B, 10B). Six observers spent approximately three hours
searching the length of Magang Cave, while three observers spent about an hour searching Shenxian cave.
FIGURE 1. Study area map and locations of caves, Fangshan World Geopark Park, Bejing, China. Parobisium magangensis sp.
n. is recorded from Magang Cave and Parobisium yuantongi sp. n. from Shenxian Cave.
TWO NEW SUBTERRANEAN-ADAPTED PSEUDOSCORPIONS Zootaxa 4661 (1) © 2019 Magnolia Press · 147
Analysis and preparation. Specimens were preserved in 75% ethanol and deposited in the Museum of He-
bei University (MHBU), Baoding City, 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. Detailed examination of characters was done using an Olym-
pus BX53 general optical microscope. Temporary slide mounts were prepared in glycerol.
Terminology. The cave environment is typically divided into four environmental zones (refer to Howarth 1980,
1983): (1) entrance zone—the combination of surface and cave environmental conditions; (2) twilight zone—both
diminished light conditions and influence of surface environment; (3) transition zone—aphotic, yet barometric
and diurnal shifts are observed at a significantly diminished rate approaching near stable climatic conditions; and,
(4) deep zone—complete darkness, high environmental stability, constant temperature, near water-saturated atmo-
sphere, and low to no airflow (usually occurs in the deepest part of the cave). The cave deep zone represents the
region most conducive to supporting subterranean-adapted animals.
Pseudoscorpion terminology and measurements mostly follow Chamberlin (1931) with some minor modifica-
tions to the terminology of the trichobothria (Harvey 1992) and chelicera (Judson 2007). The chela and chelal hand
are measured in dorsal view and all measurements are in millimeters (mm) unless noted otherwise. The following
abbreviations are 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; et = exterior terminal.
Taxonomy
Family Neobisiidae Chamberlin, 1930
Subfamily Neobisiinae Chamberlin 1930
Genus Parobisium Chamberlin 1930
Neobisium (Parobisium) Chamberlin 1930: 17; Beier 1932: 84; Morikawa 1960: 112–113; Hoff 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.
Remarks. Parobisium is characterized by the absence of a galea on the movable cheliceral finger, the fixed chelal
finger with a compact subterminal cluster of only three tactile setae (et, it, est), and a more diffuse subbasal to basal
cluster of five tactile setae (isb, ist, ib, esb, eb) (Chamberlin 1962).
Parobisium magangensis sp. n.
(Figs. 2–5)
Type material. Holotype male (Ps.-MHBU-BJ18061001): China, Beijing City, Fangshan District, Xiayunling
Town, Magang Cave, [39.711342°N, 115.745130°E], estimated cave deep zone, 751 m elevation, 10 June 2018,
Zegang Feng leg.
Paratypes: 1♂ (Ps.-MHBU-BJ18061002), 7♀ (Ps.-MHBU-BJ18061003–09), same location as holotype.
Diagnosis. Troglomorphic habitus; carapace without eyes or eyespots; epistome rounded; carapace with 6 setae
on posterior margin; pedipalp smooth and slender, both chelal fingers with 146–162 teeth; femur 8.91–8.97 times
(length 2.78–2.94), patella 7.64–7.84 times (length 2.75–2.98) longer than broad, pedicel about two-thirds of total
length of patella. Hand with pedicel 3.89–4.04 times (length 1.86–1.95), chela with pedicel 8.67–8.96 times (length
4.12–4.25) longer than broad, finger 1.18–1.23 times longer than hand with pedicel. Chelicera: rallum with 8–9 pin-
nate setae, distal one with an expanded base.
Description. Male (Fig. 3A). Carapace, chelicerae and pedipalps brown; abdomen and legs yellowish.
Carapace (Figs 4A, 5A): Smooth, 1.19–1.31 times longer than broad, with a total of 22 setae, including 4 on
anterior margin and 6 on posterior margin; lacks eyes or eyespots; epistome rounded.
FENG ET AL.
148 · Zootaxa 4661 (1) © 2019 Magnolia Press
Chelicera (Figs 4B, 5B): Hand with 6 setae, movable finger with one submedial seta; fixed finger with 13–14
teeth; movable finger with 15–16 teeth; serrula exterior with 34–37 lamellae; serrula interior with 22–25 lamellae.
Galea (Fig. 5G) replaced by a small rounded transparent spinneret. Rallum (Fig. 5C) with 8 pinnate setae, distal one
separated and with expanded base, proximal one short.
FIGURE 2. Parobisium magangensis sp. n., Female habitus.
Pedipalps (Figs 4C–D, 5H–I): Apex of coxa rounded, with 4 setae on each side. pedipalp smooth and slender.
Trochanter 3.71–3.90, femur 8.91–8.97, patella 7.64–7.84, chela (with pedicel) 8.67–8.96, chela (without pedicel)
7.59–7.71 times longer than wide, movable finger 1.61–1.78 times longer than hand (without pedicel). Fixed che-
lal finger with 8 trichobothria, movable finger with 4, eb and esb on lateral margin of hand; ib, ist, and isb closely
grouped at the base of the fixed finger; est situated at the subdistal of finger; est, et and it grouped together near
fingertip; b situated at base of movable finger, sb situated at one-third base of the finger, st and t at one-third distal
of movable finger, st nearer to t than to sb, the latter nearer to b than to st, the distance between b and sb longer than
that between t and st. Venom apparatus present only in fixed chelal finger, venom duct short. Fixed chelal finger
with 146–148 teeth, movable finger with 160–162 teeth.
Abdomen: Pleural membrane granulated. Tergal chaetotaxy (I–XI): 6: 7–9: 8: 8: 8–10: 8–9: 9–11: 8–11: 10–11:
10–11: 6–7; sternal chaetotaxy (IV–XI): 10: 12–13: 11: 11: 11: 11: 11: 2–4; stigmata with 4–6 setae around; anal
cone with 2 dorsal and 2 ventral setae. Male genital area (Figs 4E, 5E): sternite II with 22 scattered setae; sternite
III with 2–3 setae which occur on the intermediary and followed by 10 posterior setae.
Legs: Leg I (Figs 4G, 5J) and Leg IV (Figs 4H, 5K) typical. Tibia IV with one submedial tactile seta (TS = 0.54–
0.67), basitarsus IV with one basal tactile seta (TS = 0.14–0.16), telotarsus IV with one tactile seta (TS = 0.49–0.56).
Subterminal tarsal seta (Fig. 5F) bifurcate; arolium not divided, shorter than the slender and simple claws.
Female (paratypes) (Fig. 3B): Mostly same as holotype.
Chelicera. Hand with 6 setae, movable finger with 1 submedial seta; fixed finger with 13–14 teeth; movable
finger with 17–18 teeth; serrula exterior with 35–37 lamellae; serrula interior with 22–25 lamellae. Galea replaced
by a small rounded transparent spinneret; rallum of 8 blades, similar to that of holotype.
TWO NEW SUBTERRANEAN-ADAPTED PSEUDOSCORPIONS Zootaxa 4661 (1) © 2019 Magnolia Press · 149
Pedipalps. Trochanter 3.3–3.44, femur 7.53–8.18, patella 6.48–6.60, chela (with pedicel) 6.45–6.95, chela
(without pedicel) 5.64–6.05 times longer than wide, movable finger 2.09–2.30 times longer than hand (without
pedicel). Fixed chelal finger with 135–136 teeth, movable finger with 145–150 teeth.
Abdomen. Tergal chaetotaxy (I–XI): 6–8: 6–8: 7–9: 8–9: 8–10: 9–11: 9–11: 10–11: 10–12: 9–11: 6–9; sternal
chaetotaxy (IV–XI): 8–10: 10–13: 10–13: 11–12: 11–12: 11–13: 10–12: 2–4. Female genital area (Figs 3F, 4D):
sternite II with 2–3 setae on each side; sternite III with a row of 12 setae on the posterior margin.
FIGURE 3. Parobisium magangensis sp. n. A. Holotype male, dorsal view; B. Paratype female, dorsal view.
Measurements: (length/breadth or depth in mm; ratios for most characters in parentheses). Male (holotype and
paratypes). Body length 3.46–4.87. Carapace 1.19–1.31 (1.26–1.38/1.05–1.06). Pedipalpal trochanter 3.71–3.90
(1.21–1.26/0.34–0.31), femur 8.91–8.97 (2.78–2.94/0.31–0.33), patella 7.64–7.84 (2.75–2.98/0.36–0.38), chela
(with pedicel) 8.67–8.96 (4.12–4.25/0.46–0.49), chela (without pedicel) 7.59–7.71 (3.55–3.72/0.46–0.49), hand
length (without pedicel) 1.29–1.43, movable finger length 2.29–2.30 (1.61–1.78 times longer than hand without
pedicel). Leg I: trochanter 1.60–1.68 (0.40–0.42/0.25), femur 6.81–7.89 (1.43–1.50/0.19–0.21), patella 5.06–5.21
(0.91–0.99/0.18–0.19), tibia 9.23–9.57 (1.20–1.34/0.13–0.14), basitarsus 5.36–5.50 (0.59–0.66/0.11–0.12), telotar-
sus 7.27–7.70 (0.77–0.80/0.10–0.11). Leg IV: trochanter 2.88–2.92 (0.70–0.72/0.24–0.25), femur + patella 7.39–
7.81 (2.03–2.29/0.26–0.31), tibia 9.94–11.10 (1.79–2.33/0.18–0.21), basitarsus 4.43–5.07 (0.62–0.76/0.14–0.15),
telotarsus 7.15–7.42 (0.89–0.93/0.12–0.13).
Female (paratypes). Body length 4.32–5.88. Carapace 1.16–1.33 (1.41–1.45/1.06–1.25). Pedipalpal trochanter
3.36–3.44 (1.24–1.31/0.36–0.39), femur 7.53–8.18 (2.70–2.86/0.33–0.38), patella 6.48–6.60 (2.64–2.72/0.40–0.42),
chela (with pedicel) 6.45–6.95 (4.13–3.96/0.57–0.64), chela (without pedicel) 5.64–6.05 (3.45–3.61/0.57–0.64),
hand length (without pedicel) 1.35–1.39, movable finger length 2.09–2.30 (1.50–1.70 times longer than hand with-
out pedicel). Leg I: trochanter 1.43–1.58 (0.40–0.41/0.26–0.28), femur 6.24–6.68 (1.31–1.47/0.21–0.22), patella
4.94–5.17 (0.84–0.93/0.17–0.18), tibia 8.86–9.00 (1.17–1.24/0.13–0.14), basitarsus 4.67–5.27 (0.56–0.58/0.11–
0.12), telotarsus 6.00–6.55 (0.72/0.11–0.12). Leg IV: trochanter 2.58–2.59 (0.62–0.75/0.24–0.29), femur + patella
7.21–7.31 (2.09–2.34/0.29–0.32), tibia 9.76–10.85 (2.05–2.17/0.20–0.21), basitarsus 4.79–4.87 (0.67–0.73/0.14–
0.15), telotarsus 6.14–6.71 (0.86–0.94/0.14).
Distribution. This species is currently known only from the type locality.
Etymology. Latinized adjective derived from the type locality for this species, Magang Cave.
Remarks. P. magangensis sp. n. resembles Parobisium longipalpus Hong, 1996 but is distinguished by the lack
of eyes or eyespots (P. longipalpus has four conspicuous eyes), chelal fingers with 146–162 teeth (about 84–99 teeth
in P. longipalpus), and a slender pedipalpal femur 8.91–8.97 longer than broad (3.6–4.9 times in P. longipalpus),
atella 7.64–7.84 times longer than broad (2.9–3.3 times in P. longipalpus).
FENG ET AL.
150 · Zootaxa 4661 (1) © 2019 Magnolia Press
FIGURE 4. Parobisium magangensis sp. n., holotype male: A. Carapace (dorsal view); B. Right chelicerae (dorsal view); C.
Right chela (lateral view); D. Right pedipalp (dorsal view); E. Genital area of male; F. Genital area of female; G. Right leg I
(lateral view); H. Right leg IV (lateral view).
TWO NEW SUBTERRANEAN-ADAPTED PSEUDOSCORPIONS Zootaxa 4661 (1) © 2019 Magnolia Press · 151
FIGURE 5. Parobisium magangensis sp. n., holotype male (A–C, E–K), female (D). A. Carapace, dorsal view; B. Right chelic-
era, dorsal view; C. Rallum; D. Genital area of female; E. Genital area of male; F. Subterminal tarsal seta; G. Movable finger of
chelicerae, showing galea; H. Right pedipalp, dorsal view (trochanter, femur and patella); I. Right chelal, lateral view; J. Right
leg I, lateral view; K. Right leg IV, lateral view. Scale bars: 0.1 mm (C–E), 0.25 mm (B, F), 0.5 mm (A, I, J), 1 mm (G, H).
FENG ET AL.
152 · Zootaxa 4661 (1) © 2019 Magnolia Press
P. magangensis sp. n. differs from Parobisium robustiellum Hong, 1996 by lack of eyes or eyespots (P. ro-
bustiellum has four eyes), chelal fingers with 146–162 teeth (about 51–66 teeth in P. robustiellum), and a slender
pedipalpal femur 8.91–8.97 longer than broad (2.5–3.6 times in P. robustiellum), patella 7.64–7.84 times longer than
broad (2.7–3.4 times in P. robustiellum).
P. magangensis sp. n. can be easily distinguished from Parobisium anagamidense (Morikawa, 1957) by the fol-
lowing characters: carapace without eyes or eye spots (P. anagamidense with four reduced eyes), epistome rounded
(absent in P. anagamidense); chelal fingers with 146–162 teeth (about 86–90 teeth in P. anagamidense); pedipalpal
femur 8.91–8.97 longer than broad (4.0 times in P. anagamidense), patella 7.64–7.84 times longer than broad (2.9
times in P. anagamidense).
Parobisium yuantongi sp. n.
(Figs 6–8)
Type material. Holotype male (Ps.-MHBU-BJ18061010): China, Beijing City, Fangshan District, Shijiaying Town,
Shenxian cave, [39.867793°N, 115.704571°E], estimated cave deep zone, 603 m elevation, 10 June 2018, Zegang
Feng leg.
Diagnosis. Troglomorphic habitus; carapace without eyes or eyespots; epistome triangular, with rounded top;
carapace with 6 setae on posterior margin; pedipalp slender and with granulation present on femur, inside lateral of
patella and chelal hand, both chelal fingers with 115–118 teeth; femur 6.75 times (length 1.62), patella 5.70 times
(length 1.54) longer than broad, pedicel about half of total length of patella. Hand with pedicel 3.00 times (length
1.05), chela with pedicel 8.00 times (length 2.80) longer than broad, finger 1.69 times longer than hand with pedicel.
Chelicera: rallum with 8 pinnate setae, distal one with an expanded base.
Description. Male (Fig. 6). Carapace, chelicerae and pedipalps light yellow brown; abdomen and legs yellow-
ish.
FIGURE 6. Parobisium yuantongi sp. n. Holotype male, dorsal view.
TWO NEW SUBTERRANEAN-ADAPTED PSEUDOSCORPIONS Zootaxa 4661 (1) © 2019 Magnolia Press · 153
Carapace (Figs 7A, 8A): Smooth, 1.46 times longer than broad, with a total of 22 setae, including 4 on anterior
margin and 6 on posterior margin; lacks eyes or eyespots; epistome small, triangular, with rounded top.
Chelicera (Figs 7B, 8B): Hand with 6 setae, movable finger with one submedial seta; fixed finger with 14 teeth;
movable finger with 11 teeth; serrula exterior with 24–26 lamellae; serrula interior with 26–28 lamellae. Galea (Fig.
8D) replaced by a small rounded transparent spinneret. Rallum (Fig. 8C) with 8 pinnate setae, distal one separated
and with expanded base, proximal one short.
Pedipalps (Figs 7C–D, 8G–H): Apex of coxa rounded, with 4–5 setae on each side. Pedipalp slender and with
granulation present on femur, inside lateral of patella and chelal hand. Trochanter 3.33, femur 6.75, patella 5.70,
chela (with pedicel) 8.00, chela (without pedicel) 7.37 times longer than wide, movable finger 2.19 times longer
than hand (without pedicel). Fixed chelal finger with 8 trichobothria, movable finger with 4, eb and esb on lateral
margin of hand; ib, ist, and isb closely grouped at the base of the fixed finger; est situated at the subdistal of finger;
est, et and it grouped together near fingertip; b situated at base of movable finger, sb situated at one-third base of
the finger, st and t at one-third distal of movable finger, st nearer to t than to sb, the latter distinctly nearer to b than
to st, the distance between b and sb longer than that between t and st. Venom apparatus present only in fixed chelal
finger, venom duct short. Fixed chelal finger with 116 teeth, movable finger with 118 teeth.
FIGURE 7. Parobisium yuantongi sp. n., holotype male: A. Carapace (dorsal view); B. Right chelicerae (dorsal view); C. Left
chela (lateral view); D. Left pedipalp (dorsal view); E. Right leg I (lateral view); F. Right leg IV (lateral view).
FENG ET AL.
154 · Zootaxa 4661 (1) © 2019 Magnolia Press
FIGURE 8. Parobisium yuantongi sp. n., holotype male. A. Carapace, dorsal view; B. Right chelicera, dorsal view; C. Rallum;
D. Movable finger of chelicerae, showing galea; E. Subterminal tarsal seta; F. Genital area of male; G. Right chelal, lateral view;
H. Right pedipalp, dorsal view (trochanter, femur and patella); I. Right leg I, lateral view; J. Right leg IV, lateral view. Scale bars:
0.1 mm (F, G), 0.25 mm (I), 0.5 mm (B, D, E), 1 mm (A, H, J).
TWO NEW SUBTERRANEAN-ADAPTED PSEUDOSCORPIONS Zootaxa 4661 (1) © 2019 Magnolia Press · 155
Abdomen: Pleural membrane granulated. Tergal chaetotaxy (I–XI): 6: 9: 11: 11: 11: 12: 11: 11: 11: 10: 7; sternal
chaetotaxy (IV–XI): 16: 14: 14: 13: 14: 12: 11: 3; stigmata with 3–5 setae around; anal cone with 2 dorsal and 2
ventral setae. Male genital area (Fig. 8F): sternite III with 11 setae on the posterior margin; with 6 setae around the
anteromedian groove; sternite III with a row of 11 setae on the posterior margin.
Legs: Leg I (Figs 7E, 8I) and Leg IV (Figs 7F, 8J) typical. Tibia IV with one submedial tactile seta (TS = 0.50),
basitarsus IV with one basal tactile seta (TS = 0.11), telotarsus IV with one tactile seta (TS = 0.51). Subterminal
tarsal seta (Fig. 8E) bifurcate; arolium not divided, shorter than the slender and simple claws.
Measurements: (length/breadth or depth in mm; ratios for most characters in parentheses). Male. Body length
2.98. Carapace 1.46 (1.04/0.71). Pedipalpal trochanter 3.33 (0.80/0.24), femur 6.75 (1.62/0.24), patella 5.70
(1.54/0.27), chela (with pedicel) 8.00 (2.80/0.35), chela (without pedicel) 7.37 (2.58/0.35), hand length (without
pedicel) 0.81, movable finger length 1.77 (2.19 times longer than hand without pedicel). Leg I: trochanter 1.42
(0.27/0.19), femur 5.80 (0.87/0.15), patella 4.15 (0.75/0.10), tibia 7.50 (0.75/0.10), basitarsus 5.00 (0.40/0.08), telo-
tarsus 6.86 (0.48/0.07). Leg IV: trochanter 2.67 (0.46/0.18), femur + patella 5.07 (1.37/0.27), tibia 8.60 (1.29/0.15),
basitarsus 4.00 (0.44/0.11), telotarsus 6.78 (0.61/0.09).
Distribution. This species is known only from the type locality.
FIGURE 9. Magang Cave, type locality of Parobisium magangensis sp. n. and cave animals living inside: A. entrance; B.
inside the cave, showing the places where the subterranean-adapted pseudoscorpions were collected; C. Pholcus sp. (Araneae:
Pholcidae); D. Skleroprotopus membranipedalis Zhang, 1985 (Diplopoda: Julida).
Etymology. The species name, yuantongi, was derived from the Latinized Mandarin phrase for “shaped like a
cylinder” or “cylindrical.”Yuán tǒng (圆筒) refers to the shape of chelal hand.
Remarks. The new species resembles P. longipalpus Hong, 1996 but is distinguished by the lack of eyes or
eyespots (P. longipalpus has four conspicuous eyes), the slender pedipalpal femur 6.75 longer than broad (3.6–4.9
times in P. longipalpus, patella 5.70 times longer than broad (2.9–3.3 times in P. longipalpus).
P. yuantongi sp. n. also resembles P. robustiellum Hong, 1996, but it can be differentiated from the latter by
lack of eyes or eyespots (P. robustiellum has four eyes), chelal fingers with 115–118 teeth (about 51–66 teeth in P.
FENG ET AL.
156 · Zootaxa 4661 (1) © 2019 Magnolia Press
robustiellum); the slender pedipalpal femur 6.75 longer than broad (2.5–3.6 times in P. robustiellum), patella 5.70
times longer than broad (2.7–3.4 times in P. robustiellum).
P. yuantongi sp. n. can be easily distinguished from P. anagamidense (Morikawa, 1957) by the following char-
acters: carapace without eyes/eyespots (P. anagamidense (Morikawa, 1957), epistome triangular, with rounded top
(absent in P. anagamidense); pedipalpal femur 6.75 longer than broad (4.0–4.2 times in P. anagamidense (Mori-
kawa, 1957) patella 5.70 longer than broad (2.9 times in P. anagamidense (Morikawa, 1957).
P. yuantongi sp. n. differs from P. magangensis sp. n. by the pedipalpal morphology and size: pedipalp with
granulation present on femur, inside lateral of patella and chelal hand (smooth in P. magangensis sp. n.), pedipalpal
femur 6.75 longer than broad (8.91–8.97 times in P. magangensis sp. n.), patella 5.70 longer than broad (7.64–7.84
times in P. magangensis sp. n.).
FIGURE 10. Shenxian Cave, type locality of Parobisium yuantongi sp. n.: A. Surrounding surface habitat with cave entrance
(white arrow) and pig enclosure downslope from the cave entrance (red arrow) illustrated; B. deep zone environment in the cave
where P. yuantongi was collected.
Discussion
Our knowledge of cave biota in northern China is limited. Historically, speleological research has been conducted
in the subtropical or tropical southern extent of the country. However, we demonstrate in our paper that there are
also troglomorphic species in northern China that should receive equal attention from cave ecologists and resource
managers. Importantly, this work increased the number of troglomorphic pseudoscorpions in China from 20 to 22
and provides new data on the diversity of subterranean-adapted pseudoscorpions in the karst of northern China.
Both species are tentatively considered single-cave endemics. However, this may be due to limited surveys for
troglomorphic taxa rather than actual restricted distributions of these species. There are at least two caves within
a radius of 5 km in the Magang cave, and at least three caves in the radius of 5 km in the Shenxian cave. As with
many subterranean-adapted taxa (e.g., Culver et al., 2000; Christman et al., 2005; Kováč et al. 2005; Borges et al.
TWO NEW SUBTERRANEAN-ADAPTED PSEUDOSCORPIONS Zootaxa 4661 (1) © 2019 Magnolia Press · 157
2012), these two pseudoscorpion species may indeed have restricted distributions. However, they may have wider
distributions than a single cave and are likely restricted to a geologic formation rather than the caves in which they
were found. Additional surveys will be required to better define their distributional ranges, as well as access their
susceptibility to human disturbance.
Despite the impressive biological diversity found throughout China, there are no government regulations, nor
is any government agency responsible for managing and protecting cave resources. Development projects progress
steadily in karstic regions and tourist caves are developed without consideration for subterranean resources and the
rich biodiversity they often support (Whitten 2009). While we recognize environmental conditions are improving
and environmental regulations have strengthened, deforestation (Trajano 2000, Ferreira & Horta 2001, Clements et
al. 2006, Stone & Howarth 2007), intensive agriculture and water diversion (van Beynen & Townsend 2005, Stone
& Howarth 2007, Harley et al. 2011), alien species introductions (Taylor et al. 2003, Price 2004, Howarth et al.
2007), heavy metals and agrochemicals (Whitten 2009), and global climate change (Chevaldonné & Lejeune 2003,
Mammola et al. 2018) continue to present conservation challenges for cave fauna in China. Thus, we recommend
monitoring environmental conditions of the surface and subsurface of caves, as well as developing systematic in-
ventories and vulnerability assessment procedures (Mammola et al. 2019) to help safeguard China’s subterranean
natural resources.
While these longer-term programs may take more time to develop, cave biological resources in Fangshan World
Geopark are under immediate threat by human activities, and community-based conservation measures should also
be explored and potentially implemented. Shenxian cave is approximately 50 m from agriculture and human activi-
ties, and we observed direct evidence of recent human visitation to this cave. While Magang Cave is much further
from human activities (1 km), this cave was also recently visited by humans.
Gao et al. (2018) proposed several community-based conservation measures for caves in the Guangxi Province,
which may also be applicable in the north. These measures include outreach activities to educate villagers, school
children and tourists concerning the fragility of cave biological resources, as well as posting educational signs dis-
cussing the sensitivity of these resources both within the villages and potentially at proximity to the cave entrances.
Cave gating may also be appropriate in some situations, but should be endeavored only as a last resort.
Finally, to both gain inference into the distributions of these two new species, as well as to identify other poten-
tial management species, we recommend conducting additional biological inventories within these and other known
caves of the Fangshan Geopark to obtain a more comprehensive picture of northern China cave arthropod biological
diversity. We suggest applying a systematic sampling framework similar to the approach elucidated by Wynne et
al. (2018); if this is not possible, minimally researchers should aim to conduct additional direct intuitive searches in
cave deep zones, as well as bait sampling in cave deep zones to more thoroughly inventory subterranean-adapted
taxa (sensu Wynne et al. 2019).
As biological inventories continue to expand in northern China, we anticipate more subterranean-adapted ar-
thropod species will ultimately be discovered. These findings will not only better characterize the cave arthropods
of the north, but will further contribute to the already impressively rich cave fauna known to China as a whole.
Acknowledgments
We are grateful to Weitong Wang, Zhaoyi Li, Hui Wang and Yang Chen for their help in the specimens collection.
We thank also Dr. Zhizhong Gao and Dr. Xiangbo Guo for their valuable suggestions and sincere help in improv-
ing this article. Dr. Weixin Liu verified the millipede species in Figure 9. This work was supported by the National
Natural Science Foundation of China (No. 31872198), and the Ministry of Science and Technology of the People’s
Republic of China (MOST Grant No. 2015FY210300).
References
Beier, M. (1932) Pseudoscorpionidea I. Subord. Chthoniinea et Neobisiinea. Tierreich, 57, i–xx + 1–258.
https://doi.org/10.1515/9783111435107
Borges, P.A.V., Cardoso, P., Amorim, I.R., Pereira, F., Constância, J.P., Nunes, J.C., Barcelos, P., Costa, P., Gabriel, R. & Dap-
kevicius, M.L. (2012) Volcanic caves, priorities for conserving the Azorean endemic troglobiont species. International
FENG ET AL.
158 · Zootaxa 4661 (1) © 2019 Magnolia Press
Journal of Speleology, 41, 101–112.
https://doi.org/10.5038/1827-806X.41.1.11
Cao Y., Wang, Q.Q., Li, L.J. & Zhang, Y.C. (2014) Discussion on Characteristics of the Karst Cave in Beijing Fangshan District.
Urban Geology, 9 (01), 17–20. [in Chinese with English abstract]
https://doi.org/10.3969/j.issn.1007-1903.2014.01.005
Chamberlin, J.C. (1930) A synoptic classification of the false scorpions or chela-spinners, with a report on a cosmopolitan col-
lection of the same. Part II. The Diplosphyronida (Arachnida-Chelonethida). Annals and Magazine of Natural History,
Seris 10, 5, 1–48.
https://doi.org/10.1080/00222933008673104
Chamberlin, J.C. (1931) The arachnid order Chelonethida. Stanford University Publications. Biological Sciences, 7, 1–284.
Chamberlin, J.C. (1962) New and little-known false scorpions, principally from caves, belonging to the families Chthoniidae
and Neobisiidae (Arachnida, Chelonethida). Bulletin of the American Museum of Natural History, 123, 303–352.
Chamberlin, J.C. & Malcolm, D.R. (1960) The occurrence of false scorpions in caves with special reference to cavernicolous
adaptation and to cave species in the North American fauna (Arachnida-Chelonethida). American Midland Naturalist, 64,
105–115.
https://doi.org/10.2307/2422895
Christman, M.C., Culver, D.C., Madden, M.K. & White, D. (2005) Patterns of endemism of the eastern North American cave
fauna. Journal of Biogeography, 32, 1442–1452.
https://doi.org/10.1111/j.1365-2699.2005.01263.x
Chevaldonné, P. & Lejeune, C. (2003) Regional warming-induced species shift in northwest Mediterranean marine caves. Ecol-
ogy Letters, 6, 371–379.
https://doi.org/10.1046/j.1461-0248.2003.00439.x
Clements, R., Sodhi, N.S., Schilthuizen, M. & Ng, P.K. (2006) Limestone karsts of Southeast Asia: imperiled arks of biodiver-
sity. Bioscience, 56, 733–742.
https://doi.org/10.1641/0006-3568(2006)56[733:LKOSAI]2.0.CO;2
Culver, D.C., Master, L.L., Christman, M.C. & Hobbs, H.H. III. (2000) Obligate cave fauna of the 48 contiguous United States.
Conservation Biology, 14, 386–401.
https://doi.org/10.1046/j.1523-1739.2000.99026.x
Cokendolpher, J.C. & Krejca, J.K. (2010) A new cavernicolous Paribisium Chamberlin 1930 (Pseudoscorpiones: Neobisiidae)
from Yosemite National Park, U.S.A., Museum of Texas Tech University, 297,1–25.
https://doi.org/10.5962/bhl.title.156953
Ferreira, R.L. & Horta, L.C.S. (2001) Natural and human impacts on invertebrate communities in Brazilian caves. Revista
Brasileira de Biologia, 61, 7–17.
https://doi.org/10.1590/S0034-71082001000100003
Gao, Z.Z., Chen, H.M. & Zhang, F. (2017) Description of two new cave-dwelling Bisetocreagris species (Pseudoscorpiones:
Neobisiidae) from China. Turkish Journal of Zoology, 41, 615–623.
https://doi.org/10.3906/zoo-1602-39
Gao, Z.Z., Wynne, J.J. & Zhang, F. (2018) Two new species of cave-adapted pseudoscorpions (Pseudoscorpiones: Neobisiidae,
Chthoniidae) from Guangxi, China. Journal of Arachnology, 46, 345–354.
https://doi.org/10.1636/JoA-S-17-063.1
Guo, X.B & Zhang, F. (2016) Two new species of the genus Parobisium Chamberlin, 1930 from China (Pseudoscorpiones:
Neobisiidae). Entomologica Fennica, 27, 140–148.
Harley, G.L., Polk, J.S., North, L.A. & Reeder, P.P. (2011) Application of a cave inventory system to stimulate development of
management strategies: The case of west-central Florida, USA. Journal of Environmental Management, 92, 2547–2557.
https://doi.org/10.1016/j.jenvman.2011.05.020
Harvey, M.S. (1991) Catalogue of the Pseudoscorpionida. Manchester University Press, Manchester, xxx + 850 pp.
Harvey, M.S. (1992) The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Tax-
onomy, 6, 1373–1435.
https://doi.org/10.1071/IT9921373
Harvey, M.S. (2013) Pseudoscorpions of the World. Version 3.0. Western Australian Museum, Perth. Available from: http://mu-
seum.wa.gov.au/catalogues-beta/pseudoscorpions (accessed 12 October 2018)
Hoff, C.C. (1961) Pseudoscorpions from Colorado. Bulletin of the American Museum of Natural History, 122, 409–464.
Hong, Y. (1996) Two new species of the genus Parobisium (Pseudoscorpionida: Neobisiidae) from Korea. Korean Journal of
Systematic Zoology, 12, 189–197.
Howarth, F.G. (1980) The zoogeography of specialized cave animals: A bioclimatic model. Evolution, 34, 394–406.
https://doi.org/10.1111/j.1558-5646.1980.tb04827.x
Howarth, F.G. (1983) Ecology of cave arthropods. Annual Review of Entomology, 28, 365–389.
https://doi.org/10.1146/annurev.en.28.010183.002053
Howarth, F.G., James, S.A., McDowell, W., Preston, D.J. & Imada, C.T. (2007) Identification of roots in lava tube caves us-
ing molecular techniques: Implications for conservation of cave arthropod faunas. Journal of Insect Conservation, 11,
251–261. https://doi.org/10.1007/s10841-006-9040-y
TWO NEW SUBTERRANEAN-ADAPTED PSEUDOSCORPIONS Zootaxa 4661 (1) © 2019 Magnolia Press · 159
Judson, M.L.I. (2007) A new and endangered species of the pseudoscorpion genus Lagynochthonius from a cave in Vietnam,
with notes on chelal morphology and the composition of the Tyrannochthoniini (Arachnida, Chelonethi, Chthoniidae).
Zootaxa, 1627 (1), 53–68.
https://doi.org/10.11646/zootaxa.1627.1.4
Kováč, Ľ.U., Mock, A.N., Ľuptáčik, P.E., Košel, V.L., Fenďa, P.E., Svatoň, J. & Mašán, P. (2005) Terrestrial arthropods of the
Domica Cave system and the Ardovská Cave (Slovak Karst)—Principal microhabitats and diversity. 7th Central European
Workshop on Soil Zoology, April 14–16, 2003. In: Contributions to Soil Zoology in Central Europe I. Institute of Soil Biol-
ogy Academy of Sciences of the Czech Republic, České Budějovice, pp. 61–70.
Li, Y.C., Shi, A.M. & Liu, H. (2017) A new cave-dwelling species of Bisetocreagris (Arachnida, Pseudoscorpiones: Neobisi-
idae) from Yunnan Province, China. Entomology Fennica, 28, 212–218.
Liu, W. & Wynne, J.J. (2019) Cave millipede biodiversity and descriptions of eight new species from northern Guangxi, China.
Subterranean Biology, 30, 57–94.
https://doi.org/10.3897/subtbiol.30.35559
Lu, J.B. (2007) Karst Features of FangshanWorld Geopark. City Geology, 2007 (03), 26–30. [in Chinese with English abstract]
https://doi.org/10.3969/j.issn.1007-1903.2007.03.006
Lu, J.B., Lu, Y.R., Zheng, G.S. & Zheng, M.C. (2010) Formation of karst cave system and its relationship with neotectonic
movement in Beijing Western Hills, Beijing, China. Geological Bulletin of China, 29 (4), 502–509.
https://doi.org/10.3969/j.issn.1671-2552.2010.04.003
Mahnert, V. (2003) Four new species of pseudoscorpions (Arachnida, Pseudoscorpiones: Neobisiidae, Chernetidae) from caves
in Yunnan Province, China. Revue Suisse de Zoologie, 110, 739–748.
https://doi.org/10.5962/bhl.part.80209
Mahnert, V. (2009) New species of pseudoscorpions (Arachnida, Pseudoscorpiones, Chthoniidae, Chernetidae) from caves in
China. Revue Suisse de Zoologie, 116, 185–201.
https://doi.org/10.5962/bhl.part.79492
Mahnert, V. & Li, Y.C. (2016) Cave-inhabiting Neobisiidae (Arachnida: Pseudoscorpiones) from China, with description of four
new species of Bisetocreagris Ćurčić. Revue Suisse de Zoologie, 123, 259–268.
Mammola, S., Goodacre, S.L. & Isaia, M. (2018) Climate change may drive cave spiders to extinction. Ecography, 41, 233–
243.
https://doi.org/10.1111/ecog.02902
Mammola, S., Cardoso, P., Culver, D.C., Deharveng, L., Ferreira, R.L., Fišer, C., Galassi, D.P.M., Griebler, C., Halse, S., Hum-
phreys, W.F., Isaia, M., Malard, F., Martinez, A., Moldovan, O.T., Niemiller, M.L., Pavlek, M., Reboleira, A.S.P.S., Souza-
Silva, M., Teeling, E.C., Wynne, J.J. & Zagmajster, M. (2019) Scientists’ warning on the conservation of subterranean
ecosystems. BioScience, 69 (8), 641–650.
https://doi.org/10.1093/biosci/biz064
Morikawa, K. (1957) Cave pseudoscorpions of Japan (II). Memoirs of Ehime University, 2B (2), 357–365.
Morikawa, K. (1960) Systematic studies of Japanese pseudoscorpions. Memoirs of Ehime University, 2B (4), 85–172.
Price, L. (2004) An introduction to some cave fauna of Malaysia and Thailand. Acta Carsologica, 33, 311–317.
https://doi.org/10.3986/ac.v33i1.359
Ran, J.C. & Yang, W.C. (2015) A Review of Progress in Chinese Troglofauna Research. Journal of Resources and Ecology, 6,
237–246.
https://doi.org/10.5814/j.issn.1674-764x.2015.04.007
Schawaller, W. (1995) Review of the pseudoscorpion fauna of China (Arachnida: Pseudoscorpionida). Revue Suisse de Zoolo-
gie, 102, 1045–1063.
https://doi.org/10.5962/bhl.part.80489
Stone, F.D. & Howarth, F.G. (2007) Hawaiian cave biology: status of conservation and management. In: Proceedings of the
2005 National Cave and Karst Management Symposium, Albany, New York, October 31–November 4, 2005, pp. 21–26.
Taylor, S.J., Krejca, J., Smith, J.E., Block, V.R. & Hutto, F. (2003) Investigation of the potential for red imported fire ant (So-
lenopsis invicta) impacts on rare karst invertebrates at Fort Hood, Texas: A field study. In: Illinois Bexar County Karst
Invertebrates Draft Recovery Plan Natural History Survey. Center for Biodiversity Technical Report, 28, pp. 1–153.
Tian, M.Y. & Clarke, A. (2012) A new eyeless species of cave-dwelling trechine beetle from north-eastern Guizhou Province,
China (Insecta: Coleoptera: Carabidae: Trechinae). Cave and Karst Science, 39, 66–71.
Tong, Y.F. & Li, S.Q. (2008) Six new cave-dwelling species of Leptoneta (Arachnida, Araneae, Leptonetidae) from Beijing and
adjacent regions, China. Zoosystema, 30 (2), 371–386.
Trajano, E. (2000) Cave faunas in the Atlantic tropical rain forest: Composition, ecology and conservation. Biotropica, 32,
882–893.
https://doi.org/10.1111/j.1744-7429.2000.tb00626.x
van Beynen, P. & Townsend, K. (2005) A disturbance index for karst environments. Environmental Management, 36, 101–
116.
https://doi.org/10.1007/s00267-004-0265-9
Whitten, T. (2009) Applying ecology for cave management in China and neighboring countries. Journal of Applied Ecology,
46, 520–523.
FENG ET AL.
160 · Zootaxa 4661 (1) © 2019 Magnolia Press
https://doi.org/10.1111/j.1365-2664.2009.01630.x
Wynne, J.J., Sommer, S., Howarth, F.G., Dickson, B.G. & Voyles, K.D. (2018) Capturing arthropod diversity in complex cave
systems. Diversity and Distributions, 24, 1478–1491.
https://doi.org/10.1111/ddi.12772.
Wynne, J.J., Howarth, F.G., Sommer, S. & Dickson, B.G. (2019) Fifty years of cave arthropod sampling: techniques and best
practices. International Journal of Speleology, 48, 33–48.
https://doi.org/10.5038/1827-806X.48.1.2231.
Yu, W.M., Fan, W.J., Huo, S.J. & Sun, K.Q. (2011) Study on Value and Sustainable Development of Natural Heritage Sites in
North China Karst—A Case of Karst Caves in Beijing. Resource Development & Market, 27 (10), 903–906. [in Chinese
with English abstract]
https://doi.org/10.3969/j.issn.1005-8141.2011.10.013
Zhang, Y.H. & Zhu, D.H. (2012) Large karst caves distribution and development in china. Journal of Guilin University of Tech-
nology, 32 (1), 20–28. [in Chinese with English abstract]
https://doi.org/10.3969/j.issn.1674-9057.2012.01.003
... 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. 2017Tian et al. , 2018Deuve 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 identi cation of at least 21 troglomorphic pseudoscorpion species (refer to Feng et al. 2019;Li et al. 2019b). ...
... 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 identi cation 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, speci cally in Guizhou, Yunnan, Guangxi, Sichuan, and Hubei Provinces, and Beijing and Chongqing Municipalities, total at least 29 pseudoscorpion species (Schawaller 1995;Mahnert 2003Mahnert , 2009Mahnert and Li 2016;Gao 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). ...
Article
Full-text available
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.
... Interestingly, no species of Spelaeochthonius have yet been recorded from China despite the abundance of karst habitats in this country and the ongoing discovery of troglomorphic pseudoscorpions in several provinces (Gao et al. 2018(Gao et al. , 2020Feng et al. 2019Feng et al. , 2020. Recent studies indicate that Allochthonius is diverse in forest habitats throughout China, with nine species already described (Gao et al. 2016) but, again, no subterranean species has been collected there to date and all troglobiontic species belong to the Chthoniidae (genera Lagynochthonius Beier, 1951 andTyrannochthonius Chamberlin, 1929) and the Neobisiidae (Bisetocreagris Ć určić, 1983 andParobisium Chamberlin, 1930) (Feng et al. 2020) that have no known troglobiontic species in Korean caves. ...
Article
Full-text available
Two new species of troglomorphic pseudoscorpions of the family Neobisiidae, collected from karst caves in Yunnan, China, are described: Parobisium laevigatum sp. n. and P. muchonggouense sp. n.. A key to the Parobisium species from China is also provided.
Article
Full-text available
Ever-increasing human pressures on cave biodiversity have amplified the need for systematic, repeatable, and intensive surveys of cave-dwelling arthropods to formulate evidence-based management decisions. We examined 110 papers (from 1967 to 2018) to: (i) understand how cave-dwelling invertebrates have been sampled; (ii) provide a summary of techniques most commonly applied and appropriateness of these techniques, and; (iii) make recommendations for sampling design improvement. Of the studies reviewed, over half (56) were biological inventories, 43 ecologically focused, seven were techniques papers, and four were conservation studies. Nearly one-half (48) of the papers applied systematic techniques. Few papers (24) provided enough information to repeat the study; of these, only 11 studies included cave maps. Most studies (56) used two or more techniques for sampling cave-dwelling invertebrates. Ten studies conducted ≥10 site visits per cave. The use of quantitative techniques was applied in 43 of the studies assessed. More than one-third (42) included some level of discussion on management. Future studies should employ a systematic study design, describe their methods in sufficient detail as to be repeatable, and apply multiple techniques and site visits. This level of effort and detail is required to obtain the most complete inventories, facilitate monitoring of sensitive cave arthropod populations, and make informed decisions regarding the management of cave habitats. We also identified naming inconsistencies of sampling techniques and provide recommendations towards standardization.
Article
In light of recent alarming trends in human population growth, climate change, and other environmental modifications, a "Warning to humanity" manifesto was published in BioScience in 2017. This call reiterated most of the ideas originally expressed by the Union of Concerned Scientists in 1992, including the fear that we are "pushing Earth's ecosystems beyond their capacities to support the web of life. " As subterranean biologists, we take this opportunity to emphasize the global importance and the conservation challenges associated with subterranean ecosystems. They likely represent the most widespread nonmarine environments on Earth, but specialized subterranean organisms remain among the least documented and studied. Largely overlooked in conservation policies, subterranean habitats play a critical role in the function of the web of life and provide important ecosystem services. We highlight the main threats to subterranean ecosystems and propose a set of effective actions to protect this globally important natural heritage.
Article
Lagynochthonius fragilis n. sp. is described from a limestone cave in the Hong Chong karst of Kien Giang Province, southern Vietnam, which is currently threatened by quarrying activities. This is the first record of a troglomorphic species of Lagynochthonius Beier, 1951 from continental Asia. The presence of chemosensory setae on the dorsum of the chelal palm is interpreted as a synapomorphy of the tribe Tyrannochthoniini Chamberlin, 1962. The New Zealand genus Maorichthonius Chamberlin, 1925 is transferred from the Chthoniini Daday, 1888 to the Tyrannochthoniini. The genus Tyrannochthoniella Beier, 1966, also endemic to New Zealand, is assigned to the tribe Chthoniini Daday, 1888. The genus Stygiochthonius Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001, from southern Spain, is synonymized with Paraliochthonius Beier, 1956 (n. subj. syn.). Five new combinations are proposed: Lagynochthonius ovatus Vitali-di Castri, 1984 (ex Tyrannochthonius); Paraliochthonius barrancoi (Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001) (ex Stygiochthonius); P. curvidigitatus (Mahnert, 1997) (ex Lagynochthonius); P. setiger (Mahnert, 1997) (ex Tyrannochthonius); and P. superstes (Mahnert, 1986) (ex Tyrannochthonius). A key is given to the genera of the Tyrannochthoniini. The parallel evolution in several groups of pseudoscorpions of a characteristic chelal morphology, here termed lagyniform, is discussed. New designations are proposed for the spot-sensilla of the chelal fingers. The so-called ‘sensorium’ near the tip of the fixed chelal finger of Lagynochthonius species is shown to be a modified tooth that has migrated dorsally from the dental margin. The new term rallum is introduced as a replacement for the inappropriate term ‘flagellum’, as applied to the cheliceral blades of pseudoscorpions. The term bothridial vestibulum is introduced for the internal cuticular sheath at the base of the bothridia of the trichobothria.Lagynochthonius fragilis n. sp. est décrit d’une grotte calcaire de la province de Kien Giang, au sud du Vietnam, actuellement menacée par une exploitation de carrière. Elle est la première espèce troglomorphe du genre Lagynochthonius Beier, 1951 connue de l’Asie continentale. La présence des soies chemosensorielles sur la main de la pince est interprétée comme une synapomorphie de la tribu des Tyrannochthoniini Chamberlin, 1962. Le genre néo-zélandais Maorichthonius Chamberlin, 1925 est transféré des Chthoniini Daday à la tribu des Tyrannochthoniini. Le genre Tyrannochthoniella Beier, 1966, également endémique de la Nouvelle Zélande, est attribué à la tribu des Chthoniini Daday, 1888. Le genre Stygiochthonius Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001, du sud de l’Espagne, est mis en synonymie avec Paraliochthonius Beier, 1956 (n. syn. subj.). Cinq combinaisons nouvelles sont proposées : Lagynochthonius ovatus Vitali-di Castri, 1984 (ex Tyrannochthonius) ; Paraliochthonius barrancoi (Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001) (ex Stygiochthonius) ; P. curvidigitatus (Mahnert, 1997) (ex Lagynochthonius) ; P. setiger (Mahnert, 1997) (ex Tyrannochthonius) ; et P. superstes (Mahnert, 1986) (ex Tyrannochthonius). Une clé de détermination des genres de Tyrannochthoniini est fournite. L’évolution parallèle des facies caractéristiques de la pince, ici qualifié de “ lagyniforme ”, est évoquée chez plusieurs groupes de pseudoscorpions. Desnouveaux sigles sont proposés pour les sensilles punctiformes des doigts de la pince. Il est démontré que le “ sensorium ” à l’extrémité du doigt fixe de la pince des espèces de Lagynochthonius est une dent modifiée qui a migré dorsalement dès la marge dentale. Le terme inapproprié de “ flagelle ”, dans le sens de son application aux lames chélicèriennes des pseudoscorpions, est remplacé par rallum. Le terme nouveau vestibule trichobothriale est introduit pour la gaine cuticulaire à la base des bothridies des trichobothries.
Article
Two new troglomorphic pseudoscorpion species, Bisetocreagris maomaotou sp. nov. (Family Neobisiidae) and Tyrannochthonius chixingi sp. nov. (Family Chthoniidae) are described from one cave in the tower karst of northern Guangxi Province, China. This cave is located at close proximity to a village and an adjacent urban area. As with many caves in the South China Karst, this feature occurs at an elevation slightly above agriculture and rural activities; thus, we suggest it may be partially buffered from human activities in the lowland areas. We discuss the likelihood of narrow range endemism and provide research and conservation recommendations to guide future management of these two species.
Article
Aim: Identify the optimal combination of sampling techniques to maximize the detection of diversity of cave-dwelling arthropods. Location: Central-western New Mexico; northwestern Arizona; Rapa Nui, Chile. Methods: From 26 caves across three geographically distinct areas in the Western Hemisphere, arthropods were sampled using opportunistic collecting, timed searches, and baited pitfall trapping in all caves, and direct intuitive searches and bait sampling at select caves. To elucidate the techniques or combination of techniques for maximizing sampling completeness and efficiency, we examined our sampling results using nonmetric multidimensional scaling (NMDS), analysis of similarity (ANOSIM), Wilcoxon signed-rank tests, species richness estimators and species accumulation curves. Results: To maximize the detection of cave-dwelling arthropod species, one must apply multiple sampling techniques and specifically sample unique microhabitats. For example, by sampling cave deep zones and nutrient resource sites, we identified several undescribed cave-adapted and/or cave-restricted taxa in the southwestern United States and eight new species of presumed cave-restricted arthropods on Rapa Nui that would otherwise have been missed. Sampling techniques differed in their detection of both management concern species (e.g., newly discovered cave-adapted/restricted species, range expansions of cave-restricted species and newly confirmed alien species) and specific taxonomic groups. Spiders were detected primarily with visual search techniques (direct intuitive searches, opportunistic collecting and timed searches), while most beetles were detected using pitfall traps. Each sampling technique uniquely identified species of management concern further strengthening the importance of a multi-technique sampling approach. Main conclusions: Multiple sampling techniques were required to best characterize cave arthropod diversity. For techniques applied uniformly across all caves, each technique uniquely detected between ~40% and 67% of the total species observed. Also, sampling cave deep zones and nutrient resource sites was critical for both increasing the number of species detected and maximizing the likelihood of detecting management concern species.
Article
A new pseudoscorpion species, Bisetocreagris xiaoensis Li & Liu, sp. n., is described and illustrated from specimens collected in caves in Yanjin County, Yunnan Province, China. An identification key is provided to all known cave- dwelling representatives of the genus Bisetocreagris in the world.
Article
Two new pseudoscorpion species of Bisetocreagris Ćurčić, 1983, B. guangshanensis sp. nov. and B. gracilenta sp. nov., belonging to the family Neobisiidae, are described based on specimens collected in karst caves in Guizhou Province, China. Detailed diagnosis, descriptions, and illustrations are presented.