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Two new species of cave-adapted pseudoscorpions (Pseudoscorpiones, Chthoniidae) from Yunnan, China

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  • Xinzhou Normal University

Abstract and Figures

Two new cave-adapted pseudoscorpion species belonging to the family Chthoniidae are described: Tyrannochthonius pandus sp. nov. from Biyu Cave (Yunnan: Luxi) and Lagynochthonius laoxueyanensis sp. nov. from Laoxueyan Cave (Yunnan: Yanshan). Both of them, collected from the dark zone of caves, are highly troglomorphic species.
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Two new species of cave-adapted pseudoscorpions
(Pseudoscorpiones, Chthoniidae) from Yunnan, China
Yanmeng Hou1, Zhizhong Gao2, Feng Zhang1
1 e Key Laboratory of Zoological Systematics and Application, Institute of Life Science and Green
Development, Hebei University, Baoding, Hebei 071002, China 2Department of Biology, Xinzhou Teachers
University, Xinzhou 034000, Shanxi Province, China
Corresponding authors: Zhizhong Gao (gaozhizhong1987@126.com), Feng Zhang (dudu06042001@163.com)
Academic editor: Jana Christophoryová|Received 20 February 2022|Accepted 5 April 2022|Published 20 April 2022
http://zoobank.org/6FC8EE30-904F-4E8D-9EF9-996F81F30693
Citation: Hou Y, Gao Z, Zhang F (2022) Two new species of cave-adapted pseudoscorpions (Pseudoscorpiones,
Chthoniidae) from Yunnan, China. ZooKeys 1097: 65–83. https://doi.org/10.3897/zookeys.1097.82527
Abstract
Two new cave-adapted pseudoscorpion species belonging to the family Chthoniidae are described:
Tyrannochthonius pandus sp. nov. from Biyu Cave (Yunnan: Luxi) and Lagynochthonius laoxueyanensis
sp.nov. from Laoxueyan Cave (Yunnan: Yanshan). Both of them, collected from the dark zone of caves,
are highly troglomorphic species.
Keywords
Karst biotope, Lagynochthonius, taxonomy, troglobionts, Tyrannochthonius
Introduction
China has the largest karst biotopes in the world, with the karst area reaching 3.44
million km2, accounting for about one-third of the country’s land area, and contains
tens of thousands of karst caves, which are rich in animal resources (Zhao et al.
2015). Yunnan, located in southwest China, is one of the provinces with the most
widely distributed karst landforms (11.09 × 104 km2), especially in the eastern
portion of Yunnan (Wang 2001). So far, at least 742 cave-dwelling species have
been identied in China, and nearly 15% of them are from Yunnan (Latella 2019).
ZooKeys 1097: 65–83 (2022)
doi: 10.3897/zookeys.1097.82527
https://zookeys.pensoft.net
Copyright Yanmeng Hou 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.
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Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
66
Subterranean-adapted pseudoscorpions are one of the representative groups of cave-
dwelling arthropods. ey are usually eyeless, have elongate appendages and can be
easily found on cave walls or under rocks. To date, 33 cave-dwelling pseudoscorpion
species, representing three families (Chthoniidae, Neobisiidae, Chernetidae), have
been described from China. Among them, eight species are known from Yunnan
(Schawaller 1995; Mahnert 2003, 2009; Mahnert and Li 2016; Gao et al. 2017;
Li et al. 2017; Gao et al. 2018; Li et al. 2019; Feng et al. 2020; Gao et al. 2020;
Zhanget al. 2020).
e genus Tyrannochthonius Chamberlin, 1929 contains 145 species, with at least 52
species occurring in caves, and is distributed in all continents except Antarctica (WPC
2022). is genus can be diagnosed as follows (see Material and methods for explanation
of abbreviations): trichobothrium sb situated midway between st and b, or closer to st;
trichobothria ib and isb situated close together in a median or sub-basal position on the
dorsum of the chelal hand; chelal hand not distally constricted and the movable nger
without a complex or strongly sclerotized apodeme at the base; xed nger usually with
one large, medial acuminate spine-like seta at its base, but can be reduced or absent in
some cave-dwelling species; coxal spines generally long and present on coxae II only;
epistome pointed, triangular or rounded, inconspicuous and usually with 2 closely-
anking setae at its base (Chamberlin 1962; Muchmore 1984, 1991; Muchmore and
Chamberlin 1995; Edward and Harvey 2008). So far, nine species and one subspecies
of this genus have been described from China, of which six are exclusively known from
karst caves: T. akaelus Mahnert, 2009 from Sichuan, T. ganshuanensis Mahnert, 2009
from Sichuan and Hubei, T. antridraconis Mahnert, 2009 from Sichuan, T. chixingi
Gao, Wynne & Zhang, 2018 from Guangxi, T. harveyi Gao, Zhang & Chen, 2020 and
T. zhai Gao, Zhang & Chen, 2020 from Guizhou. All species are troglobites without
eyes (Mahnert 2009; Gao et al. 2018; Gao et al. 2020; WPC 2022).
e genus Lagynochthonius Beier, 1951 was erected by Beier (1951) as a subgenus
of Tyrannochthonius, but was later elevated to generic status by Chamberlin (1962).
e genus is diagnosed by trichobothrium sb situated midway between st and b, or
closer to st; trichobothria ib and isb situated close together in a median or sub-basal
position on the dorsum of the chelal hand; coxal spines generally long and present on
coxae II only; chelal hand distally constricted (or ask-shaped) and movable nger with
complex or strongly sclerotized apodeme at its base and the modied tooth (td) of the
xed chelal nger displaced onto the dorso-antiaxial face (Chamberlin 1962; Harvey
1989; Muchmore 1991; Judson 2007; Edward and Harvey 2008). At present, this
genus contains 55 species (seven species living in caves) distributed in Asia, Australia,
Africa and America. Eight species of this genus have been described from China, of
which only one is exclusively known from karst caves: L. bailongtanensis Li, Liu & Shi,
2019 from Yunnan (Li et al. 2019; WPC 2022).
Two new troglomorphic species of Chthoniidae have been recently found from the
karst caves survey in Yunnan in 2021. ese species are here described.
New cave-adapted pseudoscorpions from China 67
Materials and methods
e specimens examined for this study are preserved in 75% alcohol and deposited
in the Museum of Hebei University (MHBU) (Baoding, China) and the Museum
of Southwest University (MSWU) (Chongqing, China). Photographs, drawings and
measurements were taken using a Leica M205A stereo-microscope equipped with a
Leica DFC550 Camera and the Inkscape software (Ver. 1.0.2.0). Detailed examination
was carried out with an Olympus BX53 general optical microscope. Images were edited
and formatted using Adobe Photoshop 2022.
Terminology and measurements follow Chamberlin (1931) with some minor
modications to the terminology of trichobothria (Harvey 1992; Judson 2007) and
chelicera (Judson 2007). e chela and chelal hand are measured in lateral view and
others taken in dorsal view. All measurements are given in mm unless noted otherwise.
Proportions and measurements of pedipalps and carapace correspond to length/width,
those of legs to length/depth.
e following abbreviations are used in the text:
b basal trichobothrium;
sb sub-basal trichobothrium;
st sub-terminal trichobothrium;
t terminal trichobothrium trichobothrium;
ib interior basal trichobothrium;
isb interior sub-basal trichobothrium;
ist interior sub-terminal trichobothrium;
it interior terminal trichobothrium;
eb exterior basal trichobothrium;
esb exterior sub-basal trichobothrium;
est exterior sub-terminal trichobothrium;
et exterior terminal trichobothrium;
dx duplex trichobothria;
td modied tooth.
Taxonomy
Chthoniidae Daday, 1889
Tyrannochthonius Chamberlin, 1929
Type species. Chthonius terribilis With, 1906, by original designation.
Diagnosis. See Edward and Harvey (2008).
Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
68
Tyrannochthonius pandus sp. nov.
http://zoobank.org/D9B22241-9699-41ED-936C-5169456BD61A
Chinese name. 弯指暴伪蝎
Figs 2–5
Type material. (Figs 1A, 6) Holotype: C; Yunnan Province, Luxi County,
Luyuandong Village, the Ancient Alu Cave National Park of China, Biyu Cave;
24°34.01'N, 103°45.16'E; 1722 m a.s.l.; 13 Oct. 2021; Zegang Feng, Yanmeng Hou,
Lu Zhang and Liu Fu leg.; dark zone; Ps.-MHBU-HBUARA#2021-438-01. Paratype:
• 1; the same data as the holotype; Ps.-MSWU-HBUARA#2021-438-02.
Diagnosis. Moderately sized troglomorphic species with elongate appendages;
carapace without eyes or eyespots; anterior margin of carapace gently serrate, epistome
small, pointed, triangular, with 2 setae anking base; posterior margin of carapace
with 2 setae; tergites I–III with 2 setae; lacking chemosensory setae on dorsum of
chelal hand; chelal ngers distinctly curved in dorsal view, with numerous large, gently
curved, well-spaced teeth.
Etymology. e specic name is derived from the Latin word “pandus”, meaning
curved, refers to the curved chelal nger.
Description. Adult male (Figs 2, 3A, 4A–D, 5). Color: generally pale yellow,
chelicerae, pedipalps and tergites slightly darker, soft parts pale (Figs 2, 3A).
Cephalothorax (Figs 4B, 5A): carapace 1.07 times longer than broad, gently narrowed
Figure 1. Study area, general cave locations, and type locality for each species, Yunnan Province, China
A Biyu Cave, Tyrannochthonius pandus sp. nov. B Laoxueyan Cave, Lagynochthonius laoxueyanensis sp. nov.
New cave-adapted pseudoscorpions from China 69
posteriorly, surface smooth; anterior margin slightly serrate; without any traces of eyes;
epistome very pointed and small, triangular, with 2 setae anking base; with 18 setae
arranged s4s: 4: 4: 2: 2, most setae heavy, long and gently curved, anterolateral setae
much shorter than others; without furrows but with 4 lyrissures (Fig. 5A). Chaetotaxy
of coxae: P 3, I 3, II 4, III 5, IV 5; manducatory process with two acuminate distal setae,
anterior seta less than 1/2 length of medial seta; apex of coxa I with small, rounded
anteromedial process; coxae II with 10 terminally indented coxal spines on each side,
set as an oblique row, longer spines present in the middle of the row, becoming shorter
distally and proximally and incised for about half their length (Fig. 5C); intercoxal
tubercle absent; without sub-oral seta. Chelicera (Figs 4C, 5B): large, about as long as
carapace, 2.44 times longer than broad; 5 setae on hand, all setae acuminate, ventrobasal
seta shorter than others; movable nger with one medial seta. Cheliceral palm with
moderate hispid granulation dorsally. Both ngers well provided with teeth, xed
nger with 11 teeth, distal one largest; movable nger with 8 retrorse continuous small
teeth; galea completely vestigial (Fig.5B). Rallum with 8 blades, the distal one longest
and recumbent basally, with ne barbules and slightly set apart from the other blades,
latter tightly grouped and with long pinnae, some of which are subdivided (Fig. 5D).
Figure 2. Tyrannochthonius pandus sp. nov., male habitus. Photographed when it crawled on stony
naturalhabitat.
Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
70
Pedipalp (Figs 4A, 5E–H): surface of palpal segments smooth; setae generally long and
acuminate; femur 7.36, patella 2.55, chela 7.47, hand 2.80 times longer than deep;
movable nger 1.71 times longer than hand and 0.64 times longer than chela, without
large basal apodeme, only slightly sclerotized section present. Femur and dorsal hand
without tactile setae but with 1 distal lyrissure on patella (Fig. 5E). Fixed chelal nger
and hand with 8 trichobothria, movable chelal nger with 4 trichobothria, ib and isb
situated close together, submedially on dorsum of chelal hand; eb, esb and ist forming
a straight oblique row at base of xed chelal nger; it slightly distal to est, situated
subdistally; etslightly near to tip of xed nger, very close to chelal teeth; dx situated
distal to et; sbsituated closer to st than to b; b and t situated subdistally, b situated at
same level as est; t situated distal to est (Fig. 5F). Microsetae (chemosensory setae) absent
on hand and both chelal ngers. Sensilla absent. A tiny antiaxial lyrissure present at
base of xed nger (slightly distal to ist). Both chelal ngers with a row of teeth,
homodentate, spaced regularly along the margin, larger teeth present in the middle of
the row, becoming smaller distally and proximally: xed nger with 45 large, gently
curved, well-spaced teeth, without intercalary teeth; movable nger with 44 small
(slightly smaller than the teeth on xed nger), retrorse, gently curved and well-spaced
teeth (Fig. 5F). Chelal ngers distinctly curved in dorsal view (Fig. 5H). Opisthosoma:
generally typical; pleural membrane nely granulate. Tergites and sternites undivided;
setae uniseriate and acuminate. Tergal chaetotaxy I–XII: 2: 2: 2: 3: 4: 4: 4: 5: 5: 4:
T2T: 0; tergites VIII and IX each with an unpaired median seta. Sternal chaetotaxy
IV–XII: 10: 10: 9: 9: 9: 9: 8: 0: 2; sternites VI–IX with unpaired median seta. Anterior
genital operculum with 10 setae, genital opening slit-like, with 15 marginal setae on
each side (Fig. 4D). Legs (Fig. 5I, J): generally typical, long and slender. Femur of leg
I 1.92 times longer than patella and with 1 lyrissure at the base of femur; tarsus 2.50
Figure 3. Tyrannochthonius pandus sp. nov. A holotype male, dorsal view B paratype female, dorsal view.
Scale bars: 0.5 mm.
New cave-adapted pseudoscorpions from China 71
times longer than tibia. Femoropatella of leg IV 3.67 times longer than deep; tibia 6.29
times longer than deep; with basal tactile setae on both tarsal segments: basitarsus 4.00
times longer than deep (TS = 0.35), telotarsus 14.50 times longer than deep and 2.90
Figure 4. Tyrannochthonius pandus sp. nov., holotype male (A–D), paratype female (E) A chelal ngers
(lateral view) B carapace (dorsal view) C left chelicera (dorsal view) D male genital area (ventral view)
Efemale genital area (ventral view). Scale bars: 0.1 mm.
Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
72
Figure 5. Tyrannochthonius pandus sp. nov., holotype male A carapace (dorsal view) with a detail of
anterior margin B left chelicera (dorsal view) with details of teeth C coxal spines on coxae II (ventral
view) Drallum E left pedipalp (minus chela, dorsal view) F left chela (lateral view) with details of teeth
and with trichobothrial pattern (abbreviations explained in Material and methods) G nger tips of chela
(lateral view) H left chela (dorsal view) I leg I (lateral view) J leg IV (lateral view). Scale bars: 0.25 mm
(A–B, E–F, H–J); 0.10 mm (C–D, G).
New cave-adapted pseudoscorpions from China 73
times longer than basitarsus (TS = 0.33). Setae of leg I (trochanter to tibia) 3: 13: 12:
9, setae of leg IV (trochanter to basitarsus) 1: 3: 6: 9: 8. Arolium slightly shorter than
the claws, not divided; claws simple. Dimensions of male holotype (length/width or,
in the case of the legs, length/depth in mm; ratios in parentheses). Body length 1.41.
Pedipalps: trochanter 0.18/0.13 (1.38), femur 0.81/0.11 (7.36), patella 0.28/0.11
(2.55), chela 1.12/0.15 (7.47), hand length 0.42/0.15 (2.80), movable nger length
0.72. Chelicera 0.44/0.18 (2.44), movable nger length 0.24. Carapace 0.45/0.42
(1.07). Leg I: trochanter 0.13/0.11 (1.18), femur 0.46/0.06 (7.67), patella 0.24/0.05
(4.80), tibia 0.20/0.04 (5.00), tarsus 0.50/0.04 (12.50). Leg IV: trochanter 0.20/0.11
(1.82), femoropatella 0.66/0.18 (3.67), tibia 0.44/0.07 (6.29), basitarsus 0.20/0.05
(4.00), telotarsus 0.58/0.04 (14.50).
Adult female (Figs 3B, 4E). Mostly same as males. Anterior genital operculum
with 10 setae plus 7 setae on posterior margin. Body length 1.67. Pedipalps: trochanter
0.24/0.13 (1.85), femur 0.88/0.13 (6.77), patella 0.28/0.11 (2.55), chela 1.20/0.17
(7.06), hand length 0.49/0.17 (2.88), movable nger length 0.76. Chelicera 0.45/0.21
(2.14), movable nger length 0.23. Carapace 0.46/0.46 (1.00). Leg I: trochanter
0.14/0.11 (1.27), femur 0.50/0.06 (8.33), patella 0.26/0.06 (4.33), tibia 0.21/0.05
(4.20), tarsus 0.53/0.04 (13.25). Leg IV: trochanter 0.21/0.12 (1.75), femoropatella
0.70/0.19 (3.68), tibia 0.46/0.07 (6.57), basitarsus 0.21/0.06 (3.50), telotarsus
0.62/0.04 (15.50).
Remarks. Compared with the other six cave-dwelling species of the genus in
China, Tyrannochthonius pandus sp. nov. is most similar to T. ganshuanensis in having
only 2 setae on tergites I–III, the same chaetotaxy of carapace and triangular, a small
epistome, but diers in the shape of teeth on chelal ngers (large, gently curved, well-
spaced teeth, without intercalary teeth in T. pandus, but with pointed, well-spaced and
intercalary teeth in T. ganshuanensis), the relative position of the trichobothria on the
movable chelal nger (sb situated closer to st than to b in T. pandus, but sb situated
closer to b in T. ganshuanensis). Tyrannochthonius pandus sp. nov. can be easily separated
from T. akaleus by a smaller body size (1.67 vs. 2.10 mm in female), the teeth pattern
on chelal ngers (intercalary teeth absent in T. pandus, but present in T. akaleus); from
T. harveyi by the dierent setae number on the anterior and posterior margins of the
carapace (T. pandus with 6 and 2 setae, respectively, but T. harveyi with 4 and 4 setae,
respectively), the shape of the epistome (long and pointed in T. pandus, but rounded
and inconspicuous in T. harveyi), the number of rallar blades (8 in T. pandus, but 6 in
T. harveyi); and from T. zhai, T. chixingi and T. antridraconis by the number of setae
on the anterior tergites (tergites I–III with 2 setae in T. pandus, but the other three
with 4 setae). In addition, compared to the new species, T. zhai diers by the shorter
body length (1.40 vs. 1.67 mm in female) and lacking an epistome; T. chixingi and
T.antridraconis diers from the new species also by the presence of intercalary teeth on
the xed chelal nger (Mahnert 2009; Gao et al. 2018; Gao et al. 2020).
Distribution. is species is known only from the type locality, Biyu Cave
(Figs1A, 6). Biyu Cave is one of the tourist caves in the Ancient Alu Cave National
Park of China, with the entrance located in the Jilong hillside. is beautiful cave is
Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
74
a valley type karst cave, with an internal height of 2 to 5 m and a width of 1 to 30 m.
e cave has a latticed distribution. e stalactites in this cave are jasper colored, so it
is called Biyu Cave (a quote from the cave’s interior slogan). e specimens of this new
species were collected from under a stone and on a stone wall in an undeveloped area
Figure 6. Biyu Cave, type locality of Tyrannochthonius pandus sp. nov. A entrance B access to an unde-
veloped area C area where T. pandus specimens were collected. D, E beautiful cave landscapes inside the
cave F exit.
New cave-adapted pseudoscorpions from China 75
of the cave that is still in a natural condition. It is a small, dark, high humidity and
low temperature space (temperature: 11 °C, humidity: 90%), which is suitable for the
survival of the species.
Lagynochthonius Beier, 1951
Type species. Chthonius johni Redikorzev, 1922b, by original designation.
Diagnosis. See Judson (2007) and Edward and Harvey (2008).
Lagynochthonius laoxueyanensis sp. nov.
http://zoobank.org/4BD3052B-5217-4D93-B6FF-77DA99011228
Chinese name. 老穴岩拉伪蝎
Figs 7–10
Type material. (Figs 1B, 11) Holotype: C; Yunnan Province, Yanshan County,
Zhela Township, Liuzhao Village, Laoxueyan Cave; 23°39.03'N, 104°35.74'E; 1665 m
a.s.l.; 17 Oct. 2021; Zegang Feng, Yanmeng Hou, Lu Zhang and Liu Fu leg.; dark zone;
Ps.-MHBU-HBUARA#2021-445-01. Paratypes: • 2; the same data as the holotype;
Ps.-MHBU-HBUARA#2021-445-02, Ps.-MSWU-HBUARA#2021-445-03.
Diagnosis. Moderately sized troglomorphic species with elongate appendages;
carapace without eyes or eyespots; anterior margin of carapace thin, nely denticulate,
epistome pointed and small, triangular; posterior margin of carapace with two setae;
tergites I–II with two setae. Pedipalps slender, femur 8.54 times longer than broad;
chela 7.71 times longer than broad; chela ngers gently curved in dorsal view and xed
nger with a modied accessory tooth on dorso-antiaxial face (td).
Etymology. Latinized adjective derived from the name of the type locality,
Laoxueyan Cave, Yunnan Province, China.
Description. Adult male (Figs 8A, 9A–D, 10). Color: generally pale yellow,
chelicera, pedipalps and tergites slightly darker, soft parts pale (Fig. 8A). Cephalothorax
(Figs 9B, 10A): carapace 1.02 times longer than broad, gently narrowed posteriorly,
surface smooth, without furrows but with 1 small lyrissure; anterior margin thin,
nely denticulate; without any traces of eyes; epistome very pointed and small,
triangular; with 18 setae arranged s4s: 4: 4: 2: 2, most setae heavy, long and gently
curved, anterolateral setae much shorter than others (Fig.10A). Chaetotaxy of coxae:
P 3, I 3, II 4, III 5, IV 5; manducatory process with two acuminate distal setae,
anterior seta less than 1/3 length of medial seta; apex of coxa I with small, rounded
anteromedial process; coxae II with 9 terminally indented coxal spines on each side,
set as an oblique row, longer spines present in the middle of the row, becoming shorter
distally and proximally and incised for about half their length (Fig. 10C); intercoxal
tubercle absent; without sub-oral seta. Chelicera (Figs 9C,10B): large, about same
length as carapace, 2.37 times longer than broad; 5 setae on hand, all setae acuminate,
Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
76
Figure 7. Lagynochthonius laoxueyanensis sp. nov., female habitus. Photographed when it crawled on
stony natural habitat.
ventrobasal seta shorter than others; movable nger with one medial seta. Cheliceral
palm with moderate hispid granulation both ventral and dorsal side. Both ngers well
provided with teeth, xed nger with 12 teeth, distal one largest; movable nger with
10 retrorse continuous small teeth; galea weakly raised, keel-like (Fig. 10B). Rallum
with 8 blades, the distal one longest and recumbent basally, with ne barbules and
slightly set apart from the other blades, latter tightly grouped and with long pinnae,
some of which are subdivided (Fig. 10D). Pedipalp (Figs 9A, 10E–G): surface of palpal
segments smooth; chelal palm gradually constricted towards ngers; setae generally
long and acuminate; femur 8.54, patella 2.73, chela 7.71, hand 3.10 times longer
than broad; movable nger 1.45 times longer than hand and 0.58 times longer than
chela, apodeme complex of movable nger strongly sclerotized. Femur and dorsal hand
without tactile setae but with 1 lyrissure distally at patella (Fig. 10E). Fixed chelal nger
and hand with 8 trichobothria, movable chelal nger with 4 trichobothria, ib and isb
situated close together, submedially on dorsum of chelal hand; eb, esb and ist forming
a straight oblique row at base of xed chelal nger; it slightly distal to est, situated
subdistally; et slightly near to tip of xed nger, very close to chelal teeth; dx situated
New cave-adapted pseudoscorpions from China 77
distal to et; sb situated closer to st than to b; b and tsituated subdistally, b situated at
same level as est; t situated distal to est and at same level as it (Fig. 10F). Microsetae
(chemosensory setae) absent on hand and both chelal ngers. Sensilla absent but with
1 lyrissure between t and b, it and est, respectively (Fig.10G). Both chelal ngers with
a row of teeth, homodentate, spaced regularly along the margin, larger teeth present
in the middle of the row, becoming smaller distally and proximally: xed nger with
24 large, erect, well-spaced teeth, without intercalary teeth; movable nger with 10
small (slightly smaller than the teeth on xed nger), retrorse, serrated and well-spaced
teeth; xed nger also with a modied accessory tooth on dorso-antiaxial face (td) (Fig.
10F, G). Chelal ngers gently curved in dorsal view. Opisthosoma: generally typical;
pleural membrane nely granulate. Tergites and sternites undivided; setae uniseriate
and acuminate. Tergal chaetotaxy I–XII: 2: 2: 4: 4: 4: 4: 4: 5: 5: 4: T2T: 0; tergites
VIII and IX each with an unpaired median seta. Sternal chaetotaxy IV–XII: 13: 11: 9:
9: 9: 9: 9: 0: 2; sternites IV–X with unpaired median seta. Anterior genital operculum
with 9 setae, genital opening slit-like, with 15 marginal setae on each side (Fig. 9D).
Legs (Fig. 10H, I): generally typical, long and slender. Fine granulation present on
anterodorsal faces of trochanter IV and patella IV. Femur of leg I 1.88 times longer
than patella and with 1 lyrissure at the base of femur; tarsus 2.23 times longer than
tibia. Femoropatella of leg IV 3.76 times longer than deep; tibia 6.33 times longer than
deep; with tactile setae on both tarsal segments: basitarsus 3.57 times longer than deep,
with basal tactile setae (TS=0.24), telotarsus 12.80 times longer than deep and 2.56
times longer than basitarsus (TS=0.41). Setae of leg I (trochanter to tibia) 3: 11: 11:
14, setae of leg IV (trochanter to basitarsus) 2: 3: 6: 14: 10. Arolium slightly shorter
than the claws, not divided; claws simple. Dimensions of male holotype (length/
width or, in the case of the legs, length/depth in mm; ratios in parentheses). Body
Figure 8. Lagynochthonius laoxueyanensis sp. nov. A holotype male, dorsal view B paratype female, dorsal
view. Scale bars: 0.5 mm.
Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
78
length 1.78. Pedipalps: trochanter 0.29/0.16 (1.81), femur 1.11/0.13 (8.54), patella
0.41/0.15 (2.73), chela 1.62/0.21 (7.71), hand length 0.65/0.21 (3.10), movable
nger length 0.94. Chelicera 0.64/0.27 (2.37), movable nger length 0.34. Carapace
0.59/0.58 (1.02). Leg I: trochanter 0.17/0.11 (1.55), femur 0.60/0.07 (8.57), patella
Figure 9. Lagynochthonius laoxueyanensis sp. nov., holotype male (A–D), paratype female (E) A chelal
ngers (lateral view) B carapace (dorsal view) C left chelicera (dorsal view) D male genital area (ventral
view) E female genital area (ventral view). Scale bars: 0.1 mm.
New cave-adapted pseudoscorpions from China 79
Figure 10. Lagynochthonius laoxueyanensis sp. nov., holotype male A carapace (dorsal view) with a detail
of anterior margin B left chelicera (dorsal view) with details of teeth and tip of movable nger C coxal
spines on coxae II (ventral view) D rallum E left pedipalp (minus chela, dorsal view) F left chela (lateral
view) with details of teeth and with trichobothrial pattern (abbreviations explained in Material and meth-
ods) G nger tips of chela (lateral view) with detail of modied tooth H leg I without trochanter (lateral
view) I leg IV (lateral view). Scale bars: 0.25 mm (A, B, E , F, H, I); 0.10 mm (C , D, G).
Yanmeng Hou et al. / ZooKeys 1097: 65–83 (2022)
80
0.32/0.07 (4.57), tibia 0.30/0.05 (6.00), tarsus 0.67/0.04 (16.75). Leg IV: trochanter
0.21/0.14 (1.50), femoropatella 0.79/0.21 (3.76), tibia 0.57/0.09 (6.33), basitarsus
0.25/0.07 (3.57), telotarsus 0.64/0.05 (12.80).
Adult females (Figs 7, 8B, 9E). Mostly same as males. Anterior genital operculum
with 9 setae plus 10–12 setae on posterior margin. Body length 2.00–2.05. Pedipalps:
trochanter 0.30–0.32/0.17–0.18 (1.76–1.78), femur 1.17/0.14–0.15 (7.80–8.36),
patella 0.40–0.41/0.17 (2.35–2.41), chela 1.65–1.66/0.23–0.24 (6.88–7.22), hand
length 0.66–0.70/0.23–0.24 (2.75–3.04), movable nger length 0.95–0.98. Chelicera
0.68–0.70/0.29–0.30 (2.33–2.34), movable nger length 0.36–0.37. Carapace
0.62/0.61 (1.02). Leg I: trochanter 0.18–0.20/0.13 (1.38–1.54), femur 0.61/0.07
(8.71), patella 0.32–0.33/0.07 (4.57–4.71), tibia 0.30/0.06 (5.00), tarsus 0.68–
0.69/0.05 (13.60–13.80). Leg IV: trochanter 0.22–0.24/0.13–0.14 (1.69–1.71),
femoropatella 0.81–0.82/0.21 (3.86–3.90), tibia 0.60–0.61/0.09–0.10 (6.10–6.67),
basitarsus 0.26–0.27/0.06 (4.33–4.50), telotarsus 0.69/0.05 (13.80).
Remarks. Of all Lagynochthonius species found in hypogean environments around
the world, only three species, L. bailongtanensis Li, Liu & Shi, 2019 (from China),
L. typhlus Muchmore, 1991 (from Jamaica) and L. curvidigitatus Mahnert, 1997
Figure 11. Laoxueyan Cave, type locality of Lagynochthonius laoxueyanensis sp. nov. A entrance B the
only access to the deepest part of the cave C inside the cave entrance D area where L. laoxueyanensis
specimens were collected.
New cave-adapted pseudoscorpions from China 81
(from Spain), have no eyes, and are all highly troglomorphic species. Lagynochthonius
laoxueyanensis sp. nov. is most similar to L. typhlus in having only 2 setae on tergites
I–II, but the latter has intercalary teeth on the chelal ngers and a smaller body size
(1.28 vs. 2.00–2.05 mm in females). Lagynochthonius laoxueyanensis sp. nov. can be
easily separated from L.bailongtanensis by its smaller body size (L. laoxueyanensis 1.78
mm in male, 2.00–2.05 mm in females; while L. bailongtanensis is 2.55–2.92 mm
in males, 2.72–2.95 mm in females), the number of setae on the anterior tergites
(tergites I–II with 2 setae in L.laoxueyanensis, but 4 in L.bailongtanensis), the shape
of epistome (pointed and small in L. laoxueyanensis, but obtuse and inconspicuous
in L. bailongtanensis) and the number of setae on the pedipalpal coxa (3 setae in L.
laoxueyanensis, but 5 in L. bailongtanensis). Lagynochthonius laoxueyanensis sp. nov. can
be easily separated from L. curvidigitatus by the presence of a pair of curved chelal ngers
in the latter and the number of setae on tergites I–II (L. laoxueyanensis with 2 and 2
setae, respectively, but L. curvidigitatus with 3 and 4 setae, respectively) (Muchmore
1991; Mahnert 1997; Edward and Harvey 2008; Mahnert 2011; Li et al. 2019).
Distribution. is species is only known from the type locality, Laoxueyan Cave
(Figs 1B, 11) which is located about 4 km southeast of Liuzhao Village (Yanshan
County). e entrance of the cave is slit-shaped (18 m high and 4 m wide), and the
total length of the cave is 88.5 m, and the vertical height of the cave is about 30 m.
A descent access leads to the bottom of the cave. e bottom of the cave is a large
space, covered with gravel, temperature around 13 °C, humidity over 90%. All of the
specimens were collected from ground crevices near the end of the cave.
Acknowledgements
We thank the Ancient Alu Cave National Park of China sta for their support. We are
grateful to Zegang Feng, Lu Zhang and Liu Fu for their assistance in the eld, to Ms
Angela Xuanyu Lin for revising the language, and to Dr. Jana Christophoryová and
two reviewers, Dr. Mark S. Harvey and Dr. Katarína Krajčovičová, for their helpful
suggestions that greatly improved this paper. is work was supported by the National
Natural Science Foundation of China (No. 31872198), and the Natural Science
Foundation of Hebei Province (No. C2021201030).
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... The genus Lagynochthonius Beier, 1951, belonging to the family Chthoniidae, subfamily Chthoniinae Daday and tribe Tyrannochthoniini Chamberlin, was erected by Beier (1951) as a subgenus of Tyrannochthonius, and later elevated to generic status by Chamberlin (1962). At present, Lagynochthonius contains 56 species (the WPC lists 55 species, plus one unlisted species, L. laoxueyanensis Hou, Gao & Zhang, 2022) distributed in Asia (nine species have been described from China), Australia, Africa and America, of which eight are exclusively known from karst caves: Lagynochthonius guasirih (Mahnert, 1988) from Malaysia, L. mordor Harvey, 1989 from Australia, L. typhlus Muchmore, 1991 andL. cavicola Muchmore, 1991 from Jamaica, L. curvidigitatus Mahnert, 1997 from the Canary Islands (Spain), L. fragilis Judson, 2007from Vietnam, and L. bailongtanensis Li, Liu & Shi, 2019and L. laoxueyanensis Hou, Gao & Zhang, 2022. ...
... Of these eight species, except for L. typhlus, L. curvidigitatus, L. bailongtanensis and L. laoxueyanensis, the other four species retain eyes or eyespots and are weakly troglomorphic. It is worth mentioning that L. curvidigitatus can be easily separated from other new species described in this paper by its strongly curved chelal fingers (Fig. 1;Mahnert 1988;Harvey 1989;Muchmore 1991;Mahnert 1997;Judson 2007;Li et al. 2019;Harms et al. 2022;Hou et al. 2022;World Pseudoscorpiones Catalog 2022). ...
... Scientific research on cave-dwelling animals was first carried out in China in the early 20th century. So far, nearly 750 cave-dwelling animals have been identified in China (nearly 15% of them are from Yunnan), including 36 cave-dwelling pseudoscorpion species (ten of them are from Yunnan) (Hou et al. 2022;Latella 2019;World Pseudoscorpiones Catalog 2022;Xu et al. 2022). ...
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