Access to this full-text is provided by Pensoft Publishers.
Content available from Subterranean Biology
This content is subject to copyright. Terms and conditions apply.
Cave-dwelling millipedes, Guangxi, China 57
Cave millipede diversity with the description
of six new species from Guangxi, China
Weixin Liu1, J. Judson Wynne2
1 Department of Entomology, College of Agriculture, South China Agricultural University, 483 Wushanlu,
Guangzhou 510642, China 2 Department of Biological Sciences, Merriam-Powell Center for Environmental
Research, Northern Arizona University, Box 5640, Flagsta, AZ 86011, USA
Corresponding author: J. Judson Wynne (jut.wynne@nau.edu); Weixin Liu (da2000wei@163.com)
Academic editor: O. Moldovan | Received 19 April 2019 | Accepted 3 June 2019 | Published 02 July2019
http://zoobank.org/19205CA6-4680-45D4-A3A0-F1640752D41B
Citation: Liu WX, Wynne JJ (2019) Cave millipede diversity with the description of six new species from Guangxi,
China. Subterranean Biology 30: 57–94. https://doi.org/10.3897/subtbiol.30.35559
Abstract
We synthesized the current knowledge of cave-dwelling millipede diversity from Guangxi Zhuang Au-
tonomous Region (Guangxi), South China Karst, China and described six new millipede species from
four caves from the Guilin area, northeastern Guangxi. Fifty-two cave-dwelling millipedes are known
for the region consisting of 38 troglobionts and 14 troglophiles. Of the troglobionts, 24 are presently
considered single-cave endemics. New species described here include Hyleoglomeris rukouqu sp. nov. and
Hyleoglomeris xuxiakei sp. nov. (Family Glomeridae), Hylomus yuani sp. nov. (Family Paradoxosomati-
dae), Eutrichodesmus jianjia sp. nov. (Family Haplodesmidae), Trichopeltis liangfengdong sp. nov. (Family
Cryptodesmidae), and Glyphiulus maocun sp. nov. (Family Cambalopsidae). Our work also resulted in
range expansions of Pacidesmus tridus Golovatch & Georoy, 2014, Blingulus sinicus Zhang & Li, 1981
and Glyphiulus melanoporus Mauriès & Nguyen Duy-Jacquemin, 1997. As with many hypogean animals
in Southeast Asia, intensive human activities threaten the persistence of both cave habitats and species. We
provide both assessments on the newly described species’ distributions and recommendations for future
research and conservation eorts.
Keywords
Single cave endemic, disturbance relict, cave conservation
Subterranean Biology 30: 57–94 (2019)
doi: 10.3897/subtbiol.30.35559
http://subtbiol.pensoft.net
Copyright Weixin Liu, J. Judson Wynne . 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
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
58
Introduction
Southeast Asia is considered the most biologically diverse for troglomorphic animals
among the well-sampled tropical regions of the globe (Clements et al. 2006). Yet despite
research conducted over the past three decades, few areas in Southeast Asia have been
suciently investigated and knowledge of cave biological diversity and ecological pro-
cesses is limited (Deharveng and Bedos 2000). e expansive South China Karst rep-
resents one of these regions (Clarke 2006, Chen et al. 2001). Among the four admin-
istrative units in China where this formation occurs, Guangxi is considered the most
taxonomically well-studied region. At least 100 troglomorphic (subterranean-adapted)
arthropods have been identied from Guangxi (Wynne, unpublished data). Most have
been identied as short-range endemic species (Tian 2011, Deharveng et al. 2008) with
more than half (or 58 species) considered single cave endemics (Gao et al. 2018).
Common in cave-arthropod communities, millipedes represent a dominant and
widespread group in southern China (Golovatch 2015, Liu et al. 2017a). Approxi-
mately 150 cave-dwelling millipedes have been described from China (Golovatch
2015, Liu unpublished data) with 52 species known from Guangxi. Of these, 38 spe-
cies are troglobionts with 24 identied as single-cave endemics (Table 1). Fig. 1 sum-
marizes millipede diversity by family with Cambalopsidae (14 species), Glomeridae
(11) and Paradoxosomatidae (10) being among the most speciose groups.
In the paper, we describe six new species (representing ve families, Glomeridae,
Paradoxosomatidae, Haplodesmidae, Cryptodesmidae and Cambalopsidae), as well as
discuss range expansions of three additional cave-dwelling species. We have also sum-
marized the cave-dwelling millipedes known to Guangxi, their known distributions,
and provide recommendations to guide future research and management eorts.
Material and methods
Study area
Located in southwest China, Guangxi encompasses 236,700 km2. Once an ancient
shallow sea during the middle Cambrian to Late Triassic periods, this region is now
largely characterized by a massive karst (limestone) stratum over 10,000 m thick (Cao
et al. 2007) with steep-sided mountains called fenglin or “tower karst” protruding sky-
ward. As a result of the subtropical climate and rock stratum, Guangxi supports at least
564 known caves (Yuanhai Zhang, pers. comm. 2019).
We sampled four caves in the northeastern most extent of Guangxi within a 30 km
radius of the city of Guilin, China (Fig. 2). Caves were selected based upon two cri-
teria – sucient length to support deep zone conditions, and the availability of a cur-
rent cave map. Cave deep zones are dened as completely dark region with relatively
stable temperature, low to no airow, and a near water saturated atmosphere with a
Cave-dwelling millipedes, Guangxi, China 59
Figure 1. Species richness (total number of species provided in parentheses) for the eight known families
of subterranean-dwelling millipedes from Guangxi, China.
negligible evaporation rate (Howarth 1980, 1982). While we recognize other factors
may contribute to the occurrence of cave deep zones (e.g., mazy and/or constricted
passageways, small or partially rock-fall obstructed entrances, and cave structure in
general), we used this criterion because logistics prevented us from selecting study sites
based upon site visit evaluations. is work represents the rst eort to sample these
caves for subterranean-adapted arthropods.
All caves occurred at low elevations within tower karst formations. While extensive
agriculture, as well as rural village and suburban habitation characterized the surround-
ing lowlands, vegetation on the tower karst represented a marginally disturbed combi-
nation of native and introduced plant and tree species.
Literature review and synthesis
Based on our knowledge of the South China Karst (SCK) cave-dwelling millipede
literature, we summarized the total number of caves sampled and cave-dwelling mil-
lipedes detected for Guangxi. For selected species occurring within two or more caves,
we determined maximum distances between caves and, in some cases, average dis-
tances across all caves using ArcGIS 10.6.1.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
60
Figure 2. Study area with general locations of the four study caves (black triangles). Locator map depicts
the Guilin area (red dot), Guangxi (bold black outline), China.
Field sampling
We hand collected cave-dwelling arthropods at four caves from 15 to 18 November
2016. Approximately eight hours (2 observers at 4 hours per observer) was spent con-
ducting direct intuitive searches within estimated deep zones of each cave. We ex-
amined bat guano, dead insects, mud banks and oors, ood detritus, or vegetation
Cave-dwelling millipedes, Guangxi, China 61
brought in by humans and deposited within the cave. Given the four caves varied in
size and the diversity of arthropods encountered, the areal extent sampled varied. We
also sampled the vegetation within the entrance of Liangfeng Cave applying a direct
intuitive search approach. For all four caves, we also opportunistically collected arthro-
pods as encountered while transiting from entrance to estimated deep zones. Sampling
methods were applied sensu stricto Wynne et al. (2019).
Analysis and preparation
All specimens examined in this study were collected by the second author in the Guilin
area, Guangxi, southern China and preserved in 95% ethanol. Holotypes and para-
types are deposited in the Zoological Collection of the South China Agricultural Uni-
versity, Guangzhou, Guangdong Province, China (SCAU). Detailed examination of
characters and dissections were performed using a Leica S8 APO stereo micro scope.
Line drawings were prepared with a ZEISS Axioskop40 microscope with a camera lu-
cida attached. Photographs were taken using a Keyence VHX-5000 digital mi croscope,
and further edited using Adobe Photoshop CS5.
Cave locations
We recognize standard practice for new species description is to provide sample locality
information including geographical data to facilitate future collecting, interpretation
and research. Because caves often represent sensitive resources, we provided general
geographical information and oset the latitude and longitude coordinates by ~1 km.
is level of detail is sucient for future comparative studies, while protecting the
precise location of our study caves.
Terminology
Cave ecosystems typically consist of four zonal environments (Howarth 1980, 1983):
(1) entrance zone – combination of surface and cave environmental conditions; (2)
twilight zone – both diminished light conditions and inuence of surface environ-
ment; (3) transition zone – aphotic, yet barometric and diurnal shifts are observed at a
signicantly diminished rate approaching near stable climatic conditions; and, (4) deep
zone – complete darkness, high environmental stability, constant temperature, and
near water-saturated atmosphere with low to no airow (typically occurs in the deepest
portion of the cave). While there are four primary cave specic functional groups gen-
erally recognized, the specimens discussed are troglomorphic (subterranean-adapted)
organisms known as troglobionts (Sket 2008). ese animals are obligate cave dwellers
that require the stable environmental conditions of the deep zone to complete their life
cycle and exhibit morphological characteristics indicative of cave adaptation. We also
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
62
Table 1. Fifty-two known cave-dwelling millipede species from Guangxi, South China Karst, China. ‘Or-
der: Family: Species’, ‘Functional Group’ (TB=presumed troglobiont, TP=troglophile), number of caves
(# Caves) to suggest a level of potential endemism), and main Reference(s) are provided. As troglophiles
are expected to have regional distributions, the number of caves where TPs were detected is not included.
Order: Family: Species Functional group # Caves Reference(s)
Glomerida: Glomeridae
Hyleoglomeris baxian Liu & Tian, 2015 TB 1 Liu and Tian 2015a
Hyleoglomeris curtisulcata Golovatch, Liu & Georoy, 2012 TB 1 Golovatch et al. 2012a
Hyleoglomeris grandis Liu & Tian, 2015 TB 1 Liu and Tian 2015a
Hyleoglomeris heshang Golovatch, Liu & Georoy, 2012 TB 1 Golovatch et al. 2012a
Hyleoglomeris kunnan Golovatch, Liu & Georoy, 2012 TB 1 Golovatch et al. 2012a
Hyleoglomeris lii Golovatch, Liu & Georoy, 2012 TP – Golovatch et al. 2012a
Hyleoglomeris mashanorum Golovatch, Liu & Georoy,
2012
TB 1 Golovatch et al. 2012a
Hyleoglomeris mulunensis Golovatch, Liu & Georoy, 2012 TB 1 Golovatch et al. 2012a
Hyleoglomeris rukouqu sp. nov. TP – is study
Hyleoglomeris xueju Golovatch, Liu & Georoy, 2012 TB 1 Golovatch et al. 2012a
Hyleoglomeris xuxiakei sp. nov. TB 1 is study
Polydesmida: Cryptodesmidae
Trichopeltis liangfengdong sp. nov. TB 1 is study
Haplodesmidae
Eutrichodesmus distinctus Golovatch, Georoy, Mauriès &
VandenSpiegel, 2009
TB 1 Golovatch et al. 2009a
Eutrichodesmus jianjia sp. nov. TB 1 is study
Eutrichodesmus latus Golovatch, Georoy, Mauriès &
VandenSpiegel, 2009
TB 4 Golovatch et al. 2009b
Eutrichodesmus lipsae Golovatch, Georoy, Mauriès &
VandenSpiegel, 2015
TB 1 Golovatch et al. 2015
Eutrichodesmus planatus Liu & Tian, 2013 TB 1 Liu and Tian 2013
Eutrichodesmus similis Golovatch, Georoy, Mauriès &
VandenSpiegel, 2009
TB 2 Golovatch et al. 2009b
Paradoxosomatidae
Piccola golovatchi Liu & Tian, 2015 TB 1 Liu and Tian 2015b
Hylomus longispinus (Loksa, 1960) TB 1 Loksa 1960, Golovatch et al. 2010a,
Srisonchai et al. 2018
Hylomus lui (Golovatch, Li, Liu & Georoy, 2012) TB 1 Golovatch et al. 2012b, Srisonchai et
al. 2018
Hylomus nodulosus (Liu, Golovatch & Tian, 2014) TP – Liu et al. 2014, Srisonchai et al. 2018
Hylomus phasmoides (Liu, Golovatch & Tian, 2016) TB 1 Liu et al. 2016, Srisonchai et al. 2018
Hylomus scolopendroides Golovatch, Georoy & Mauriès,
2010
TB 4 Golovatch et al. 2010a, Liu et al.
2014, Srisonchai et al. 2018
Hylomus scutigeroides Golovatch, Georoy & Mauriès,
2010
TB 6 Golovatch et al. 2010a, Liu et al.
2014, Srisonchai et al. 2018
Hylomus spinissimus (Golovatch, Li, Liu & Georoy, 2012) TB 1 Golovatch et al. 2012b, Srisonchai et
al. 2018
Hylomus variabilis (Liu, Golovatch & Tian, 2016) TP – Liu et al. 2016, Srisonchai et al. 2018
Hylomus yuani sp. nov. TB 1 is study
Polydesmidae
Epanerchodus orientalis Attems, 1901 TP – Golovatch et al. 2012c
Pacidesmus armatus Golovatch, Georoy & Mauriès, 2010 TB 3 Golovatch et al. 2010b
Pacidesmus bedosae Golovatch, Georoy & Mauriès, 2010 TB 3 Golovatch et al. 2010b
Pacidesmus bidus Golovatch & Georoy, 2014 TB 1 Golovatch and Georoy 2014
Pacidesmus tiani Golovatch, Georoy & Mauriès, 2010 TB 2 Golovatch et al. 2010b
Pacidesmus tridus Golovatch & Georoy, 2014 TB 4 Golovatch and Georoy 2014, is
study
Cave-dwelling millipedes, Guangxi, China 63
Order: Family: Species Functional group # Caves Reference(s)
Callipodida: Paracortinidae
Paracortina yinae Liu & Tian, 2015 TP – Liu and Tian 2015c
Spirostreptida: Pericambalidae
Bilingulus sinicus Zhang & Li, 1981 TP – Zhang and Li 1981, is study
Parabilingulus aramulus Zhang & Li, 1981 TP – Zhang and Li 1981
Parabilingulus simplicius Mauriès & Nguyen Duy-
Jacquemin, 1997
TP – Mauriès and Nguyen Duy-Jacquemin
1997
Cambalopsidae
Hypocambala polytricha Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB 1 Golovatch et al. 2011a
Glyphiulus acutus Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB? 2 Golovatch et al. 2011b
Glyphiulus calceus Jiang, Guo, Chen & Xie, 2018 TP – Jiang et al. 2018
Glyphiulus dicilis Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB 2 Golovatch et al. 2011b
Glyphiulus echinoides Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TP – Golovatch et al. 2011c
Glyphiulus foetidus Jiang, Guo, Chen & Xie, 2018 TP – Jiang et al. 2018
Glyphiulus impletus Jiang, Guo, Chen & Xie, 2018 TP – Jiang et al. 2018
Glyphiulus maocun sp. nov. TB 1 is study
Glyphiulus melanoporus Mauriès & Nguyen Duy-
Jacquemin, 1997
TP – Mauriès and Nguyen Duy-Jacquemin
1997, This study
Glyphiulus mulunensis Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB 2 Golovatch et al. 2011b
Glyphiulus paramulunensis Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB 2 Golovatch et al. 2011b
Glyphiulus proximus Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB 2 Golovatch et al. 2011b
Glyphiulus speobius Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB 2 Golovatch et al. 2011b
Glyphiulus tiani Golovatch, Georoy, Mauriès &
VandenSpiegel, 2011
TB 1 Golovatch et al. 2011b
reference troglophiles (or troglophilous organisms) – non-troglomorphic animals that
occur facultatively within caves and complete their life cycles there, but also exist in
similar cave-like habitats on the surface (Barr 1967, Howarth 1983, Sket 2008). Terms
used for taxonomic descriptions follow Minelli (2015).
Results
Since ~1960, at least 49 caves have been sampled for cave-dwelling millipedes. Of
these, nearly half (24 caves) supported single-cave endemic troglobionts, nine caves
supported at least two troglobitic millipede species, and three caves (present study;
Table 2) contained three troglobionts. For species occurring in two or more caves,
maximum distance between caves ranged from 11.22 km for Glyphiulus speobius to
137.27 km for Hylomus scutigeroides (Table 3).
A total of nine cave-dwelling millipede species were collected from four caves near
Guilin (Table 2). Four millipede species were detected in Liangfeng Cave, the other
three caves contained three species each. Of the six undescribed species, four are troglo-
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
64
Table 2. Nine cave-dwelling millipedes including both troglobitic and potentially relict species from four
caves near Guilin, Guangxi, South China Karst, China.
Species Maomaotou cave Guanshan No. 4 cave Shangshuiyan cave Liangfeng cave
Hyleoglomeris rukouqu sp. nov. – – × –
Hyleoglomeris xuxiakei sp. nov. – × – –
Hylomus yuani sp. nov. – – – ×
Eutrichodesmus jianjia sp. nov. – × – –
Trichopeltis liangfengdong sp. nov. – – – ×
Pacidesmus tridus × × × –
Blingulus sinicus × – × ×
Glyphiulus melanoporus × – – –
Glyphiulus maocun sp. nov. – – – ×
Table 3. Fourteen troglomorphic millipede species detected in more than one cave, Guangxi, South
China Karst, China. ‘Order: Family: Species’, number of caves (# Caves) to suggest a level of potential
endemism), and maximum distance between caves in kilometers (Max Dist.) are provided. *Indicates
coordinate data for all caves was unavailable for that species. ‘?’ denotes maximum distance was indeter-
minable due to lack of cave coordinate data. ‘Landscape features’ potentially aecting distribution of caves
separated by more than 10 km.
Order: Family: Species # Caves Max dist. (km) Landscape features
Polydesmida: Haplodesmidae
*Eutrichodesmus latus 4 14.32 Maximum distance between two caves provided, the third
cave occurs in between the two.
*Eutrichodesmus similis 2 ?
Paradoxosomatida
Hylomus scolopendroides 4 81.27 ree caves clustered within 21.24 km; one cave separated by
Diaojiang, Hongshui and Jincheng Rivers and lowland areas
Hylomus scutigeroides 6 137.6 One cave separated from others by Hongshui River; others
may occur in same formation
Polydesmidae
*Pacidesmus armatus 3 12.4 Two of three caves separated by lowland areas
*Pacidesmus bedosae 3 2.67
Pacidesmus tiani 2 1.88
Pacidesmus tridus 4 59.7 Separated by lowland areas
Spirostreptida: Cambalopsidae
Glyphiulus acutus 2 8.85
Glyphiulus dicilis 2 9.32
Glyphiulus mulunensis 2 7.77
Glyphiulus paramulunensis 2 6.67
Glyphiulus proximus 2 2.67
Glyphiulus speobius 2 11.22 Separated by lowland areas
morphic and may represent single cave and/or short-range endemics. One undescribed
epigean species, Hyleoglomeris rukouqu sp. nov., was identied within cave entrance
vegetation, and may represent a ‘disturbance relict’ species (sensu stricto Wynne et al.
2014). Glyphiulus maocun sp. nov., a troglophile, was detected in one cave, but likely
has a more regional distribution. For known troglophiles, Blingulus sinicus Zhang &
Li, 1981 were more widely distributed occurring in three caves each, while Glyphiulus
melanoporus Mauriès & Nguyen Duy-Jacquemin, 1997 was detected in one cave.
Cave-dwelling millipedes, Guangxi, China 65
Class Diplopoda
Order Glomerida Leach, 1814
Family Glomeridae Leach, 1815
Genus Hyleoglomeris Verhoe, 1910
Type species. Hyleoglomeris multilineata Verhoe, 1910.
Remarks. e genus can be characterized by the telopods, which with frontomesal tri-
chosteles on the prefemur and femur, and the caudomesal femoral process forming a dis-
tinct angle to femur proper (Golovatch et al. 2006, Liu and Tian 2015a). It encompasses
over 90 species distributed in Eurasia from Japan and Sulawesi to the Balkans (Engho et
al. 2015). At present, 30 species of this genus occur in China, 25 are subterranean-adapted.
Hyleoglomeris rukouqu sp. nov.
http://zoobank.org/EFA33915-ECD8-4B25-B874-28D0354B907C
Figs 3A, 5, 6
Type material. Holotype male (SCAU), China, Guangxi Zhuang Autonomous Re-
gion, Yangshou County, Shangshuiyan Cave [24°57'43.6"N, 110°20'37.21"E], 191
m elevation (el.), cave entrance, ferns and other vegetation (refer to habitat section
below), direct intuitive search, 17 November 2016, J.J. Wynne leg. Paratype, 1 female
(SCAU), same data as holotype.
Etymology. e species name, rukouqu, is used as a noun in apposition from the
Mandarin phrase, rùkǒu qū (入口区). When translated to English it means “entrance
zone” or “entrance area” to denote the area in the cave where this species was collected.
Diagnosis. Adult male of H. rukouqu sp. nov. is distinct from other Hyleoglomeris
species based on the following combination of characters: (1) peculiar color pattern
(Fig. 5); (2) telopods with a large, rectangular, central syncoxital lobe (Fig. 6D). is
new species is similar to H. lii (a troglophile from a cave in Guangxi), but is distin-
guished by (1) 2+2 dark brown spots on the thoracic shield (Fig. 5B, C) vs. 1+1 light
grey-yellow spots in H. lii; (2) sycoxital lobe of telopods being high, large and rectan-
gle-shaped (Fig. 6D) vs. low, linguiform, apically evidently concave in H. lii.
Description. Based on type specimens. Length ca 8.0 mm (holotype), 11.0 mm
(paratype), width 5.0 mm (holotype), 7.0 mm (paratype). Coloration: pattern vivid (Fig.
3A). With the exception of the brown dorsal spots, this species had similar coloration as
that of the limestone rock and sediment within the entrance. General coloration in alco-
hol (Fig. 5) light yellowish with contrastingly dark spots, latter absent on rings 3 and 11.
Head only brownish caudal margin, antennae and ommatidia dark-brownish. Collum
mostly marbled dark-brown except frontal margin part. oracic shield with 2+2 dark
brown spots, but lateral ones smaller than middle ones. Terga 4 with 1+1 brownish spots,
smaller than the above. Terga 5–7 and pygidium with 1+1 large, oblong, transverse, para-
median, marbled, dark spots. Terga 8–10, these paramedian spots increasingly separated
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
66
Figure 3. A Hyleoglomeris rukouqu sp. nov. from Shangshuiyan Cave B Hyleoglomeris xuxiakei sp. nov.
from Guanshan No. 4 Cave C Hylomus yuani sp. nov. from Liangfeng Cave D Eutrichodesmus jianjia sp.
nov. from Guanshan No. 4 Cave.
into 2+2, but lateral ones smaller than middle ones. Head: Ommatidium at least 7+1,
lenses rather convex. Tömösváry’s organ transverse-oval, parallel to the body, only slightly
wider than long. Antennae with four apical cones, antennomere 6 ca 2.5 times as long as
Cave-dwelling millipedes, Guangxi, China 67
Figure 4. A Trichopeltis liangfengdong sp. nov. from Liangfeng Cave B Pacidesmus tridus Golovatch &
Georoy, 2014 from Maomaotou Cave C Glyphiulus maocun sp. nov. from Liangfeng Cave.
Figure 5. Hyleoglomeris rukouqu sp. nov., holotype. A–C Habitus, dorsal, lateral and ventral views, respectively.
wide. Exoskeleton: Collum with two transverse striae (Fig. 5C). oracic shield (Fig. 5B)
with a narrow hyposchism, the latter not reaching behind caudal tergal margin; 7 trans-
verse striae: 3 starting below, 1 level to, 3 above schism; 5 striae (never the rst and last
from below) crossing the dorsum. Following terga 3–11 with two striae above lateral edge
(Fig. 5B). Pygidium (last tergite) of both sexes slightly concave medially at caudal mar-
gin. ♂ leg 17 with a low, rounded, outer coxal lobe; telopodite 4-segmented (Fig. 6A). ♂
leg 18 with a subtrianglar syncoxital notch; telopodite 4-segmented (Fig. 6B). Telopods:
(Fig. 6C–E) with a large, subrectanglar, central syncoxital lobe anked by high setose
horns, each of the latter with a small lobe on top. Prefemur micropapillate laterally, with
a long, digitiform, frontomesal trichostele. Femur with a smaller, digitiform, frontomesal
trichostele. Caudomedial femoral process prominent, apically with an evident lobe. Tibia
with a frontomesal seta. Caudomesal tibial process evident, recurved; an indistinct, papil-
late tubercle at base on caudal face. Tarsus strongly sigmoid, narrowly rounded apically.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
68
Figure 6. Hyleoglomeris rukouqu sp. nov., holotype. A leg 17, anterior view B leg 18; anterior view
Cright half of telopods, posterior view D right half of telopods, anterior views E tip of syncoxital horn,
anterior view.
Habitat. Specimens were collected within a vegetation association that may be
limited to the cave entrance zone and similar geographic features (e.g., sinkholes and
ssures in rock). At least three plant species occurred within the entrance including
Gesneriaceae sp., Adiantum sp., and one other fern species, which cannot be identied
without examining the sorii (A. Monro, pers. comm. 2018).
Notes. Based on the vivid color pattern and well-developed ommatidia, as well as
the location where it was detected within the cave, this animal represents an epigean
species and may be functioning as an obligate troglophile (sensu Peck 1970). While it
is likely H. rukouqu sp. nov. either occurred or still occurs in similar habitats on the
surface, the importance of relict plant species restricted to cave entrances has been
discussed for southern China (Monro et al. 2018). Additionally, several arthropod spe-
cies globally are restricted to cave entrances in Polynesia (Mockford and Wynne 2013,
Wynne et al. 2014, Bernard et al. 2015, Taiti and Wynne 2015) and North America
(Benedict 1979, Wynne and Shear 2016) due to either extensive surface disturbance
and glacial interglacial cycles, respectively. us, it is possible this species is a ‘distur-
bance relict’ restricted to the entrance of Shangshuiyan Cave and potentially other area
cave entrances with similar vegetation.
Hyleoglomeris xuxiakei sp. nov.
http://zoobank.org/D3CA0855-76FD-4683-8A7C-3F0505DC3D8E
Figs 3B, 7, 8
Type material. Holotype male (SCAU), China, Guangxi Zhuang Autonomous Re-
gion, Yangshuo County, Guanshan No. 4 Cave [24°56'58.34"N, 110°20'53.52"E],
Cave-dwelling millipedes, Guangxi, China 69
186 m el., deep zone, direct intuitive search, 16 November 2016, J.J. Wynne leg.
Paratype, 1 male (SCAU), same data as holotype. 3 females (SCAU), same data as
holotype, but deep zone, slightly muddy at area.
Etymology. e species name, xuxiakei, was Latinized using a combination of the
surname and forename of Xu Xiake (徐霞客). Xu was a traveler, explorer, and the rst
speleologist of China, who studied caves and karst geology during the decline of the
Ming Dynasty. He conducted a four-year expedition (1636–1640) across southern
China where he examined over 300 caves (including more than 100 from the Guilin
area; Sweeting 1995, Ravbar 2016). In his book, Xu Xiake’s Travels (rst published in
1642), he described underground streams and ponds, provided sketch maps of caves,
as well as proposed terminology for karst features; many of his speleological terms are
still used today (Ravbar 2016).
Figure 7. Hyleoglomeris xuxiakei sp. nov., holotype. A–C Habitus, dorsal, lateral and ventral views,
respectively.
Figure 8. Hyleoglomeris xuxiakei sp. nov., holotype. A Leg 17, anterior view B leg 18; anterior view
Cright half of telopods, posterior view D right half of telopods, anterior views.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
70
Diagnosis. Adult male of H. xuxiakei sp. nov. is distinct from other Hyleoglomeris
species based on the following combination of characters: (1) nearly pallid color (Fig.
7); (2) ♂ leg 17 with 3-segmented telopodite (Fig. 8A); (3) telopods with a low, oval-
shaped, central syncoxital lobe; (4) horns of syncoxital lobe without any structure on
top (Fig. 8D). is new species is clearly distinguished by the depigmented body (Fig.
3B) vs. vivid color pattern in H. rukouqu sp. nov. (Fig. 3A).
Description. Based on type specimens. Lengths of body ca 3.8–4.5 mm, width 2.5
–3.0 mm in both sexes. Coloration: entirely pallid (Figs 3B, 7). Head: Ommatidium
at least 5(6) + 1, translucent, barely visible (Fig. 7C). Tömösváry’s organ transverse-ov-
al, parallel to the body, only slightly wider than long. Antennae with four apical cones,
antennomere 6 ca 2.0 (♂) or 1.8 (♀) times as long as wide. Exoskeleton: Collum with
two transverse striae (Fig. 7C). oracic shield with a narrow hyposchism, the latter
reaching behind caudal tergal margin; 8–9 transverse striae: 4(5) starting below, one
level to, 3(4) above schism; 5 striae (never the rst and last from below) crossing the
dorsum. Following terga 3–11 with two striae above lateral edge. Pygidium of both
sexes regularly rounded at caudal margin. ♂ leg 17 with a low, subrounded, outer
coxal lobe; telopodite 3-segmented (Fig. 8A). ♂ leg 18 with an arch-shaped syncoxital
notch; telopodite 4-segmented (Fig. 8B). Telopods: (Fig. 8C–D) with a rather low,
oval-shaped, transverse, central syncoxital lobe anked by high setose horns, each of
the latter without any structure on top. Prefemur micropapillate laterally, with a well-
developed frontomesal trichostele. Femur with a smaller frontomesal trichostele. Cau-
domesal femoral process prominent, apically with an evident lobe strongly curved to
frontad. Tibia with a frontomesal seta. Caudomesal tibial process evident, recurved; a
distinct, papillate tubercle at base on caudal face. Tarsus strongly sigmoid, narrowly
rounded apically.
Habitat. is species was collected from a chamber within the estimated cave deep
zone, approximately 50 m from the cave entrance. Cave sediment was compact mud
with a small amount of rock breakdown from the ceiling.
Notes. Based on a depigmented habitus and translucent ommatidia, we consider
this species is a troglobiont.
Order Polydesmida Leach, 1815
Family Paradoxosomatidae Daday, 1889
Genus Hylomus Cook & Loomis, 1924
Type species. Hylomus draco Cook & Loomis, 1924.
Remarks. e genus is often referred to as “dragon millipedes” because of the para-
terga is antler-like, wing-shaped or spiniform. It can be characterized by the collum and
metaterga often with granulate, tuberculate and/or speculate; male femora (5, 6, 7 and/
or 9) often humped ventrally; and gonopods mostly suberect, some subfalcated, a short-
ened solenomere mostly sheathed by a usually condensed, rather simple solenophore
Cave-dwelling millipedes, Guangxi, China 71
(Liu et al. 2014, 2016). is genus encompasses 33 species distributed from China,
Laos, Vietnam and ailand (Srisonchai et al. 2018). At present, 19 species are known
in China and 13 are subterranean-adapted (Liu et al. 2016; Srisonchai et al. 2018).
Hylomus yuani sp. nov.
http://zoobank.org/89255A98-84A8-4D57-ADF4-F7B4F0BDBE48
Figs 3C, 9–12
Material examined. Holotype male (SCAU), China, Guangxi Zhuang Autonomous
Region, Lingchuan County, Liangfeng Cave [25°12'34.86"N, 110°31'56.8"E], 184 m
el., deep zone, direct intuitive search, 18 November 2016, J.J. Wynne leg. Paratypes, 1
male, 3 females (SCAU), same data as holotype.
Etymology. is species is named for Dr. Yuan Daoxian (袁道先), a preeminent
and globally recognized Chinese hydrologist. He has made numerous signicant con-
tributions in advancing both the science and management of karst and karst waters in
southern China since the 1960s.
Diagnosis. Adult males of H. yuani sp. nov. are distinct from other Hylomus spe-
cies based on the following combination of characters: (1) paraterga long and spini-
form only on collum and rings 2–6, short and coniform thereafter (Figs 9, 10A, B);
(2) collum with 8+8 frontal, 3+3 in the middle, 3+3 caudal, setigerous spinules; (3) ♂
femur 6 strongly inated in distal 1/4 (Fig. 10D); (4) gonopod with a agelliform sole-
nomere and a strongly condensed solenophore (Figs 9–12). is new species is similar
to H.lui (a troglobiont from a cave in Guilin), but is distinguished by (1) paraterga
long and spiniform on collum and rings 2–6 (Fig. 9) vs. on collum and rings 2–5 in
H. lui; (2) collum and following metaterga with three transverse rows of spinules (Fig.
10A–B) vs. two transverse rows of spinules in H. lui.
Description. Based on type specimens. Lengths of body ca 27–28 (♂), 30–35 mm
(♀); width of mid-body pro- and metazonae 1.5–1.6 and 2.0 (♂), 2.0–2.2 and 2.5–2.6
mm (♀). Coloration: light brownish to nearly pallid (Figs 3C, 9–10). Antennomere
7 dark brown. Body: with 20 rings. In width, head < ring 2 < collum < 4–5 < 3 = 6–7
< 8–9 < 10–13; thereafter body gradually tapered posteriorly towards telson. Head:
densely setose, but more sparsely on vertex, epicranial suture conspicuous (Fig. 10A).
Antennae long and slender, reaching past ring 8 (♂) or 7 (♀) when extended posteri-
orly. Exoskeleton: Collum with 8+8 evident setigerous spinules arranged in a row at
front margin, behind it with 3+3 in the middle and 3+3 similarly spinules at posterior
margin. Metaterga 2–4 each with 5+5, 3+3 and 5+5(6) similarly setigerous spinules ar-
ranged in three transverse rows (Fig. 9A). Following metaterga with same sculpture, but
7–9+7–9 setigerous spinules at posterior margin, 2–4 spinules of them extended to the
lateral side (Figs 9C, 10B). Paraterga clearly spiniform, but well-developed only in para-
terga 1–6, directed more dorsad than laterad (Fig. 9A). Following paraterga increasingly
stout and short, paraterga 7–14 directed dorsolaterad, thereafter directed clearly caudad
(Figs 9C, 10A–B). Tegument shining, prozonae delicately microalveolate, metaterga
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
72
Figure 9. Hylomus yuani sp. nov., ♂ paratype. A Anterior part of body, dorsal view B anterior part of
body, ventral view C posterior part of body, dorsal view D posterior part of body, ventral view.
Cave-dwelling millipedes, Guangxi, China 73
Figure 10. Hylomus yuani sp. nov., ♂ paratype. A Mid-body rings, dorsal view B mid-body rings, lateral
views C sternite V, ventral view D femora 6 and 7, subventral view.
and surface below paraterga nely microgranulate. Constriction between pro- and
metazonae broad and shallow (Fig. 10A–B). Pore formula normal; transverse sulcus
usually very vague, but traceable in rings 5–19 (Fig. 10A). Ozopores inconspicuous,
located near the base on lateral side of pore-bearing paraterga (Fig. 10B). Pleurosternal
carinae evident only on rings 2 and 3 in both sexes, absent thereafter. Epiproct simple,
lateral pre-apical papillae distinct (Fig. 9C). Hypoproct subtrapeziform, caudal margin
emarginate, setigerous cones at caudal edge large, widely separated (Fig. 9D). Sterna
sparsely setose, cross-shaped impressions weak. A paramedian pair of short, rounded,
independent tubercles between ♂ coxae 4 (Figs 9B, 10C). Legs long and slender, ca
5.0 (♂) or 3.5 (♀) times as long as mid-body ring height. ♂ femur 6 strongly inated
ventrally in distal 1/4 (Fig. 10D). Gonopods: (Figs 11, 12) Coxite short, subcylindrical,
densely setose distodorsally, about 1/3 as long as telopodite. Prefemur densely setose
and about half as long as acropodite. Femorite elongate, slightly curved, with seminal
groove running entirely on mesal face. Solenophore (sph) strongly condensed; soleno-
mere (sl) agelliform, evidently separated at base from solenophore.
Habitat. is species was collected within the estimated cave deep zone, approximate-
ly 20 m from the cave entrance. Cave sediment was compact mud and the passageway was
swaddled by a slightly visible fog. is species and cave crickets (family Rhaphidophoridae)
were among the most abundant species within this portion of cave deep zone.
Notes. Based on the slender elongate antennae and legs, a depigmented cuticle, the
species is considered a troglobiont.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
74
Figure 11. Hylomus yuani sp. nov., ♂ paratype. A Right gonopod, lateral view B right gonopod, mesal views.
Figure 12. Hylomus yuani sp. nov., ♂ paratype. A Right gonopod, lateral view B right gonopod, mesal
views. Designations: sl = solenomere; sph = splenophore.
Cave-dwelling millipedes, Guangxi, China 75
Family Haplodesmidae Cook, 1895
Genus Eutrichodesmus Silvestri, 1910
Type species. Eutrichodesmus demangei Silvestri, 1910.
Remarks. e genus can be characterized by the body often capable of volvation,
with or without mid-dorsal projections; paraterga 2 strongly enlarged; and gonopod
femorite with a more or less distinct process or outgrowth laterally (Golovatch et al.
2009a, b). It encompasses 53 species distributed from south Japan, southern China, and
Southeast Asia to Vanuatu, Melanesia (Engho et al. 2015, Liu et al. 2017b). At pre-
sent, 23 species have been known in continental China, 20 are subterranean-adapted.
Eutrichodesmus jianjia sp. nov.
http://zoobank.org/403323E4-C776-4DC8-8795-9C53C3B64E02
Figs 3D, 13, 14
Type material. Holotype male (SCAU), China, Guangxi Zhuang Autonomous Re-
gion, Yangshuo County, Guanshan No. 4 Cave [24°56'58.34"N, 110°20'53.52"E],
186 m el., deep zone, direct intuitive search, 16 November 2016, J. J. Wynne leg.
Paratypes, 4 females, 1 juv. (SCAU), same data as holotype; 1 male (SCAU), same data
as holotype, but collected on mud bank.
Etymology. e species name, jianjia (肩甲), is used as a noun in apposition. is
phrase was used to denote the well-developed paraterga, which covers the millipede’s
legs much like the pauldrons of body armor cover the shoulder of the warrior.
Diagnosis. Adult males of E. jianjia sp. nov. is distinct from other Eutrichodesmus
species based on the following combination of characters: (1) collum with ve, and
metaterga 2–19 with three transverse rows of round microvillose tubercles (Fig. 13).
(2) gonopod acropodite with a dorsolateral tooth (t) at midway and apical with a large
tube-shaped lobe (l) (Fig. 14). is new species is clearly distinguished from E. lipsae (a
troglobiont from a cave in Guilin) by (1) metaterga without mid-dorsal projections vs.
most metaterga with mid-dorsal projections; (2) gonopod acropodite with a triangular,
ventral process (p) at about basal 1/3 (Fig. 14) vs. without this process in E. lipsae.
Description. Based on type specimens. Lengths of adult body ca 6.0–7.0 mm,
widths of mid-body pro- and metazonae 0.6–1.0 and 1.2–1.3 mm in both sexes. Col-
oration: generally light-yellowish to pallid (Fig. 3D). Body: subcylindrical, congloba-
tion complete, adults with 20 rings (Fig. 10A). Head: frons densely setose, nely micro-
granulate on vertex, with a paramedian pair of rounded knobs above antennal sockets.
Epicranial suture conspicuous. Antennae short, slightly clavate. Exoskeleton: Collum
subtrapeziform, slightly broader than head, not covering the latter from above, with ve
transverse rows of round microvillose tubercles (Fig. 13B). Metaterga 2–19 each with
three transverse mixostictic rows of similar tubercles extending onto paraterga, about
6–8+6–8 per row. e middle row of tubercles slightly larger than others. Pre-anal ring
short, with four transverse rows of small tubercles (Fig. 13D, E). Paraterga with evident
shoulders anteriorly, strongly declivous, broad and usually triobate laterally, evidently
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
76
Figure 13. Eutrichodesmus jianjia sp. nov., ♂ paratype. A Habitus, lateral view B anterior part of body,
ventral view C anterior part of body, dorsal view D posterior part of body, ventral view E posterior part
of body, lateral view.
Figure 14. Eutrichodesmus jianjia sp. nov., ♂ paratype. A Left gonopod, lateral view B left gonopod,
mesal view. Designations: dp = distofemoral process; l = lobe; p = process; t = tooth.
extending down below level of venter; caudolaterally at base with two distinct lobula-
tions (Fig. 13A). Paraterga 2 strongly enlarged. Tergal setae and ozopores invisible. Ozo-
pores hardly visible. Prozonae nely alveolate; constriction between pro- and metazoan
Cave-dwelling millipedes, Guangxi, China 77
narrow and shallow. Epiproct apically with four spinnerets. Hypoproct subtrapeziform,
with two long setae. Sterna narrow, but much broader between ♂ coxae 6–7 and 9.
Gonopod aperture suboval. Legs long and slender, but hardly reaching tips of paraterga.
Gonopods: (Fig. 14) Coxite large, abundantly setose and micropapillate ventrolaterally.
Telopodite subfalcate, distinctly curved ventrad, setose in its basal part, with a promi-
nent, denticulate, lateral, distofemoral process (dp) at midway. Acropodite with a trian-
gular, ventral process (p) at about basal one-third and a dorsolateral tooth (t) at midway;
apical with a large tube-shaped lobe (l). Seminal groove long, terminating without hair.
Habitat. Specimens were collected in the same general location as H. xuxiakei sp.
nov. specimens. us, refer to the H. xuxiakei sp. nov. habitat description.
Notes. Based on the long slender legs and a depigmented cuticle, we consider this
species a troglobiont.
Family Cryptodesmidae Karsch, 1880
Genus Trichopeltis Pocock, 1894
Type species. Cryptodesmus bicolor Pocock, 1894.
Remarks. e genus can be characterized by the metaterga distinctly several trans-
verse rows of tuberculations, usually setose, and gonopod usually foliate, with a tri-
partite or deeply notched telopodite. It encompasses 12 species distributed from the
Himalayas of India, through Bangladesh, Myanmar to southern China, Laos, Viet-
nam, Cambodia and Indonesia (Likhitrakarn et al. 2017). At present, 5 species have
been known in China, 4 are subterranean-adapted.
Trichopeltis liang fengdong sp. nov.
http://zoobank.org/7E972E70-ADDA-4CA1-924F-3016EF5E3FA7
Figs 4A, 15–17
Type material. Holotype male (SCAU), China, Guangxi Zhuang Autonomous Re-
gion, Lingchuan County, Liangfeng Cave [25°12'34.86"N, 110°31'56.8"E], 184 m
el., deep zone, steam bank, direct intuitive search, 18 November 2016, J.J. Wynne leg.
Paratypes, 4 females (SCAU), same data as holotype.
Etymology. is species name, liangfengdong, is used as a noun in apposition and
is the name of the type locality.
Diagnosis. Adult male of T. liangfengdong sp. nov. is distinct from other Trichopeltis
species based on the following combination of characters: (1) metaterga 2–15 with four
transverse rows of small, setigerous tubercles (Fig. 15B); (2) gonopod coxae without
seta and acropodite with several small subapical lobules (los) (Fig. 17). is new species
is similar to T. reexus (a troglobiont from a cave in Hunan), but is distinguished by (1)
metaterga 2–15 with four transverse rows of setigerous tubercles vs. two transverse rows
in T. reexus; (2) tip of acropodite bid (Fig. 17) vs. distorted in T. reexus.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
78
Description. Based on type specimens. Length of both sexes ca 14.0–15.0 mm,
widths of mid-body pro- and metazonae 1.5–2.0 and 4.8–5.0 mm. Coloration: gener-
ally pallid (Fig. 4B). Body: with 20 rings (Fig. 15). In width, collum < ring 2 < 3 = 4
< 5 < 6 < 7–16; thereafter body gradually tapered posteriorly towards telson. Head:
densely pilose and microgranulate, epicranial suture present (Fig. 16A). Antennae
long, reaching past ring 3 when extended posteriorly; in length, antennomere 6 = 5 >
4 = 3 > 2 > 7 > 1. Exoskeleton: Collum fan-shaped (Fig. 15A), covering the head from
above, dorsal surface with irregular several small, round, setigerous tubercles. Marginal
lobules on collum: 13+13 small, setigerous, rounded anteriorly, 7+7 relative larger,
laterally and 5+5 very small caudally. Mid-dorsal regions on rings 2–15 with four more
or less regular, transverse rows of similarly small, setigerous tubercles, 4–5 + 4–5 per
row (Figs 15B, 16B). Several tubercles extending onto paraterga. Following metaterga
with ve rows of smaller tubercles, 6–7 + 6–7 per row (Fig. 15B). Paraterga strong-
ly developed, clearly upturned dorsally above the dorsum only on the collum, other
paraterga at. Each with 5–6 small, dentiform, lateral and 7–8 much larger, squarish
caudolateral lobules, all evident, setigerous and microvillose (Figs 15, 16A, B). Cau-
Figure 15. Trichopeltis liangfengdong sp. nov. holotype. A Habitus, ventral view B habitus, sublateral view.
Cave-dwelling millipedes, Guangxi, China 79
Figure 16. Trichopeltis liangfengdong sp. nov., holotype. A anterior part of body, ventral view B anterior part
of body, dorsal view C sterna 6 and 7, ventral view D right gonopod, mesal view E right gonopod, lateral view.
dolateral lobules on paraterga mostly oblong, relatively large, and well separated from
one another (Fig. 16A). Caudolateral corner of paraterga projecting behind rear ter-
gal margin on rings 15–19 (Fig. 15). Integument clearly microgranulate throughout,
prozonae nely alveolated. Limbus regularly crenulated. Stricture between pro- and
metazonae broad, shallow and nely microgranulated. Tergal setae simple and short
(Fig. 16A, B). Ozopores invisible, pore formula untraceable. Epiproct short, with four
spinnerets apically. Hypoproct subtrapeziform, 1+1 caudal setigerous papillae clearly
separated. Pleurosternal carinae present on rings 2 and 3 in both sexes. Sterna mod-
estly setose, cross-shaped impressions moderate, broadened between ♂ coxae 9 (Fig.
16C). Legs long and slender, unmodied, produced beyond paratergal lateral margin
(Fig. 15), about 1.8 times as long as mid-body ring height in both sexes. Gonopods:
(Figs 16D, E, 17) Coxite short and squarish, without seta. Prefemur densely setose and
a particularly long setae; nearly half the length of telopodite. Femorite strongly at-
ten, pie-shaped, with a small lobe (l) ventrally. Acropodite folded, with several small
subapical lobules (los), tip bid. Seminal groove terminating with a hairy pulvillus,
forming no distinct solenomere.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
80
Habitat. Specimens were collected in the same general location as H. yuani sp.
nov. specimens. us, refer to the H. yuani sp. nov. habitat description.
Notes. Based on the long slender antennae and legs and a depigmented cuticle, the
species is considered a troglobiont.
Family Polydesmidae Leach, 1815
Genus Pacidesmus Golovatch, 1991
Type species. Pacidesmus shelleyi Golovatch, 1991
Remarks. e genus can be characterized by metaterga often with three transverse
rows of 3+3 sculpture, and gonopod structure showing no prominent clivus to recurve
laterad of the seminal groove (Golovatch and Georoy 2006). It encompasses 9 spe-
cies, only the type species from a mountain in Chiengmai, ailand, all other 8 species
have been known from caves in China.
Figure 17. Trichopeltis liangfengdong sp. nov., holotype. A Left gonopod, mesal view B left gonopod,
lateral view. Designations: l = lobe; los = lobules.
Cave-dwelling millipedes, Guangxi, China 81
Pacidesmus tridus Golovatch & Georoy, 2014
Fig. 4B
Material examined. 6 males, 3 females (SCAU), China, Guangxi Zhuang Autonomous
Region, Yangshuo County, Guanshan No. 4 Cave [24°56'58.34"N, 110°20'53.52"E],
186 m el., deep zone, direct intuitive search, 16 November 2016, J.J. Wynne leg. 1
male, 1 juv. (SCAU), China, Guangxi Zhuang Autonomous Region, Xiufeng District,
Maomaotou Cave [25°18'46.12"N, 110°16'12.64"E], 225 m el., deep zone, direct in-
tuitive search, 15 November 2016, J. J. Wynne leg. 2 males, 1 female, 4 juv. (SCAU),
China, Guangxi Zhuang Autonomous Region, Yangshou County, Shangshuiyan Cave
[24°57'43.6"N, 110°20'37.21"E], 191 m el., deep zone, direct intuitive search, 17
November 2016, J. J. Wynne leg.
Notes. is troglobiont was rst found in Skeleton Cave [25°17'13"N, 110°13'26"E],
el. 186 m (Golovatch and Georoy 2014). We later conrmed P. tridus within three caves
with the furthest cave (Guanshan # 4) approximately ~13 km to the south of type locality.
e three caves where we detected P. tridus are separated by a distance of ~41 km. e
northernmost cave, Maomaotou Cave occurs 39.6 km north from Guangshan No. 4 Cave
and Shangshuiyan Cave, which are co-located occurring within 1 km of each other. As it
is currently morphologically described as one species, it now has a regional distribution.
Habitat. is species was collected from the estimated deep zones of the three
caves. For Maomaotou cave, we collected it within bamboo detritus at mid-cave. In
Shangshuiyan Cave, specimens were collected within decomposing detritus and along
muddy cave sediment deposits within a sinuous passageway. For Guangshan No. 4
Cave, refer to habitat descriptions of H. xuxiakei sp. nov.
Order Spirostreptida Brandt, 1833
Family Pericambalidae Silvestri, 1909
Genus Bilingulus Zhang & Li, 1981
Type species. Bilingulus sinicus Zhang & Li, 1981.
Remarks. e genus can be characterized by the lingular lamellae divided into two
parts. ♂ legs 1 with a syncoxite carry a pair of long, band-like coxal process; femur
with a nger-shaped process at the end protruding posteriorly, tarsus without claw at
the end. Anterior gonopod with a very broad coxa; telopodite of posterior gonopod
slender, devoid of branch (Zhang and Li 1981).
Bilingulus sinicus Zhang & Li, 1981
Material examined. 3 males, 3 females, 5 juv. (SCAU), China, Guangxi Zhuang
Autonomous Region, Yangshou County, Shangshuiyan Cave [24°57'43.6"N,
110°20'37.21"E], 191 m el., deep zone, cartography station #19, direct intuitive
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
82
search, 17 November 2016, J.J. Wynne leg. 4 females (SCAU), China, Guangxi
Zhuang Autonomous Region, Xiufeng District, Maomaotou Cave [25°18'46.12"N,
110°16'12.64"E], 225 m el., deep zone, direct intuitive search, 15 November 2016,
J.J. Wynne leg. 1 juv. (SCAU), China, Guangxi Zhuang Autonomous Region, Ling-
chuan County, Liangfeng Cave [25°12'34.86"N, 110°31'56.8"E], 184 m el., deep
zone, direct intuitive search, 18 November 16, J.J. Wynne leg.
Notes. is species lacked characters suggestive of troglomorphy. It had a pigment-
ed cuticle and ommatidia were present. us, B. sinicus is considered a troglophile. It
was rst described by Zhang and Li (1981) from a cave in Guilin; no cave name or
coordinate data were provided. While we detected this species in three dierent Guilin
caves, we’re unable to determine a distance from the type locality. e caves where we
detected P. tridus is separated by an average distance of 34.1 km. Maomaotou Cave
occurs to the north-northwest and is 45 km from Liangfeng Cave and 40.4 km to the
north of Shangshuiyan Cave. Based upon its epigean characteristics and that this spe-
cies is regionally distributed on the landscape, we consider it a troglophile.
Family Cambalopsidae Cook, 1895
Genus Glyphiulus Gervais, 1847
Type species. lulus (recte: Julus) granulatus Gervais, 1847.
Remarks. e genus has been divided into two groups, granulatus-group and ja-
vanicus-group. Both groups shared the same gonopods characters as the anterior gono-
pods usually with a plate-like coxosternum, 1-segmented telopodite. Posterior gonop-
ods highly compressed, showing a plumose, subagelliform, distal process (Golovatch
et al. 2007). It encompasses over 60 species distributed in Southeast Asia and southern
China. At present, 42 species have been known in China, 34 are subterranean-adapted.
e below two species belong to granulatus-group because ♂ legs 1 strongly reduced,
sternum devoid of median structures.
Glyphiulus melanoporus Mauriès & Nguyen Duy-Jacquemin, 1997
Material examined. 1 male, 1 female, 1 juv. (SCAU), China, Guangxi Zhuang Autono-
mous Region, Xiufeng District, Maomaotou Cave, [25°18'46.12"N, 110°16'12.64"E],
225 m el., deep zone, direct intuitive search, 15 November 2016, J. J. Wynne leg.
Notes. G. melanoporus is considered a troglophile. e type locality for this species
is Skeleton Cave [25°17'13"N, 110°13'26"E] (Mauriès and Nguyen Duy-Jacquemin
1997). We collected this species from Maomaotou Cave, which represents a 21.1 km
range expansion to the south of the type locality.
Habitat. For our specimens, we collected them within the deep zone of Maomao-
tou Cave. Sediment where specimens were collected was compacted and wet.
Cave-dwelling millipedes, Guangxi, China 83
Glyphiulus maocun sp. nov.
http://zoobank.org/5CDE1D3A-71A2-48EA-B4BB-F9FDD20D860D
Figs 4C, 18–20
Type material. Holotype male, China, Guangxi Zhuang Autonomous Region, Ling-
chuan County, Maocun Village, Liangfeng Cave [25°12'34.86"N, 110°31'56.8"E],
184 m el., deep zone, direct intuitive search, 18 November 2016, J.J. Wynne. Para-
types, 4 females, 4 juv. (SCAU), same data as holotype.
Etymology. e species name, maocun, is used as a noun in apposition and is the
name of the village nearest to the type locality.
Diagnosis. is new species is similar to G. melanoporus (from caves in Guilin)
in showing carinotaxy formula of collum I–IV+5c+6a+pc+ma. However, G. maocun
sp. nov. is distinct from G. melanoporus by (1) carinotaxy formula of metaterga 2/2+I/
Figure 18. Glyphiulus maocun sp. nov., holotype. A , B Anterior part of body, sublateral views C mid-
body rings, dorsal view D posterior part of body, ventral view.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
84
Figure 19. Glyphiulus maocun sp. nov., holotype. A Head and collum, dorsal view B head, collum, and
legs 1, ventral view C legs 1, anterior view D leg 2, caudal view E legs 3, frontal view F anterior gonopods,
caudal view G posterior gonopods, anterior view H posterior gonopods, caudal view.
i+3/3; (2) anterior gonopod with a conspicuous, high, curved downward process on
coxosternum (Figs 19F, 20A); (3) posterior gonopod tip branch plumose agellum
(Figs 19G–I, 20B, C).
Cave-dwelling millipedes, Guangxi, China 85
Figure 20. Glyphiulus maocun sp. nov., holotype. A Anterior gonopods, caudal view B posterior gonop-
ods, anterior view C posterior gonopods, caudal view.
Description. Based on the type specimens. Lengths of both sexes ca 26–32 mm,
mid-body rings round in cross-section, their widths and heights similar, 1.2–1.8 mm.
Coloration: yellow-brown to brownish in alcohol. Ommatidia blackish (Fig. 18A).
Body: with 46–54 podous rings + 2–1 apodous one(s) + telson. Head: clypeus with
4 teeth anteromedially. Each eye patch with about 8–12 ommatidium arranged in
three irregular vertical rows (Figs 18A, 19A). Antennae long, antennomere 7 with
four short apical cones (Fig. 18A, B). Gnathochilarium usually, with a separate pro-
mentum (Fig.19B). Exoskeleton: Carinotaxy formula of collum I–IV+5c+6a+pc+ma,
anterior margin crests lower (Fig. 19A). Subsequent metaterga strongly crested, cari-
notaxy formula 2/2+I/i+3/3. Ozoporiferous tubercles round (Fig. 18). Prozonae deli-
cately alveolate; strictures and metazonae ne longitudinal striations. Rings 2 and 3
with long pleural aps. Epiproct simple, with a short, low, rounded tubercle medially.
Paraprocts rather regularly convex. Hypoproct broadly emarginated (Fig. 18D). ♂
legs 1 very strongly reduced, represented only by a sternum devoid of any median or
paramedian structures but carrying 1+1 strongly separated prongs both curved ante-
riad and bearing several strong setae (Fig. 18B, C). ♂ legs 2 with large coxae; penes
oblong, each with two strong setae distolaterally (Fig. 19D). ♂ legs 3 with slender
and elongate coxae (Fig. 19E). Legs slender, about 1.2 times as long as mid-body ring
height. Claw simple, without any spine at base. Anterior gonopods: (Figs 19F, 20A)
with a conspicuous, high, lobe-shaped, curved downward process on coxosternum,
which is much higher than telopdite; telopodite slightly coiled, 1-segmented, lateral
in position, with 2 or 3 strong apical setae. Posterior gonopods: (Figs 19G–I, 20B, C)
laterally with a low, lateral lobe carrying a dozen of long setae, tip branch very slender
and high, plumose agellum.
Habitat. Specimens were collected within or at proximity to ood detritus.
Notes. Specimens exhibited no characteristics suggestive of cave adaptation. It had
a pigmented cuticle and well-developed blackish ommatidia. We consider this species
to be troglophillic within Liangfeng Cave. Subsequently, this species may have a larger,
more regional distribution.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
86
Discussion
Our work increased the number of subterranean-adapted millipedes from 34 to 38
species. With 564 known caves in Guangxi (Yuanhai Zhang, pers. comm. 2019), but
a fraction (8.7% or 49 caves) of these caves have been examined for troglomorphic
millipedes. e identication of six new species of millipedes from four caves in the
Guilin area underscores the potential for many additional discoveries. For millipede-
specic surveys, direct intuitive searches of selected cave deep zones (sensu Wynne et
al. 2019) has been identied as the best technique; Mesibov et al. (1995) reported that
four weeks of hand collecting during the fall was more ecient than an intensive pitfall
trapping eort involving trapping seven days per month for over one year.
Conversely, while advancing our knowledge of regional troglomorphic millipede
diversity stands to contribute signicantly to the conservation value of the SCK, a more
comprehensive understanding (i.e., for all cave-dwelling taxa) of regional diversity, as
well as advancing procedures to assessing the vulnerability of cave systems to anthro-
pogenic impact should be considered. Wynne et al. (2018, 2019) provides a system-
atic framework and guidance for sampling cave-dwelling arthropods using a repeatable
framework, which can be modied for application in the SCK. Using such an approach,
these data may be then be examined by applying principles similar to the cave vulner-
ability assessment developed by Tanalgo et al. (2018). rough such an eort, system-
atic techniques may be used to assemble robust and comparable landscape scale data,
and each cave subsequently be evaluated for its vulnerability due to human activities.
Moreover, for conservation biologists and resource managers to best evaluate the
importance of SCK cave biological diversity, we will require a more robust understand-
ing of the distributional ranges of troglomorphic species (as well as other subterranean-
restricted taxa). Specically, while nearly 40 percent of troglomorphic millipedes were
identied as single cave endemics, 14 troglobionts (including one whose range was
expanded from this work) occurred within two or more caves. One species, P. tridus,
was conrmed within three caves in the Guilin region (two caves from this work, plus
the type locality); the maximum distance between caves for this species was 59.7 km.
ree additional subterranean-adapted species occurred in multiple caves with maxi-
mum distances ranging from 81.27 and 137.6 km (Table 3). As many troglomorphic
arthropods are identied as short-range endemic species, occurring in a single cave
or geological formation (Reddell 1994, Culver et al. 2000, Christman et al. 2005,
Deharveng et al. 2008, Tian 2011, Harvey and Wynne 2014, Gao et al. 2018, Nitzu
et al. 2018) and that rivers and valleys/ lowland areas often result in vicariance (Barr
1985, Faille et al. 2015, Katz et al. 2018), the genetic relatedness of at least these three
species should be further examined using genetic techniques. While these species may
be morphological similar, we suggest they may be genetically distinct – potentially
representing dierent subspecies or lineages.
We also reported a possible ‘disturbance relict’, H. rukouqu sp. nov., discovered
within a cave entrance vegetation community. As similar cave entrance vegetation
Cave-dwelling millipedes, Guangxi, China 87
communities have been identied as either supporting distinct relict plant communi-
ties and/or plant species in southern China (Monro et al. 2018), Easter Island, Chile
(Wynne et al. 2014), and west-central New Mexico, USA (Lindsey 1951, Northup and
Welbourn 1997, Wynne 2013), their importance in supporting cave-restricted arthro-
pod populations demonstrated (Northup and Welbourn 1997, Wynne 2013, Wynne
et al. 2014, Wynne and Shear 2016), and widespread land cover conversion of both
lowlands and uplands has occurred in China since 1958, this nding warrants both
additional research into this species distribution, as well as a larger scale examination
of other potential ‘disturbance relict’ arthropod species within cave entrance vegetation
communities of the SCK.
Although caves are often considered distinct from the surface environment, cave
ecosystems are inextricably linked to surface processes. Caves require an allochthonous
energy supply, which may include ood detritus, guano deposition from bats, birds and
crickets, and dissolved organic materials that percolate from the surface. us, when
humans adversely change the surface environment, the cave ecosystem may change as
well. Deforestation (Trajano 2000, Ferreira and Horta 2001, Clements et al. 2006,
Stone and Howarth 2007), intensive agriculture and water diversion (van Beynen
and Townsend 2005, Stone and Howarth 2007, Harley et al. 2011), livestock grazing
(Stone and Howarth 2007), alien species introductions (Elliott 1992, Reeves 1999,
Taylor et al. 2003, Howarth et al. 2007, Price 2016), heavy metals and agrochemicals
(Whitten 2009), and global climate change (Chevaldonné and Lejeune 2003, Mam-
mola et al. 2018) have negatively aected both cave organisms and ecological processes.
Despite the impressive biological diversity found in China, there are no govern-
ment regulations, nor is any government agency responsible for managing and protect-
ing cave resources. Development projects typically progress in caverniferous regions
and tourist caves are developed without consideration for subterranean resources and
the rich biodiversity they often support (Whitten 2009). While we recognize environ-
mental conditions are improving and environmental regulations have strengthened in
China, Whitten (2009) and Cao et al. (2007) identied most of the aforementioned
impacts continue to expand and severely stress both epigean and hypogean ecosystems
in southern extent of the country. us, we recommend monitoring environmental
conditions of the surface and subsurface of caves identied as supporting rare endemic
species populations and/or sensitive ecosystems.
is paper revealed that at least 38 troglomorphic millipede species occur in
Guangxi. Other examples of SCK’s high cave diversity includes at least 19 species of
troglomorphic pseudoscorpions (Gao et al. 2018), 29 vascular plant species believed
restricted to cave entrances (Monro et al. 2018), and the richest globally diversity of
caveshes with at least 154 species and nearly half of these species considered single
cave endemics (Zhao et al. in press). As this work continues, we anticipate the number
of subterranean-adapted millipedes, as well as other arthropod species will increase.
rough ours and other eorts, we believe the SCK will ultimately emerge as a global
hotspot for cave biological diversity.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
88
Acknowledgements
Dr. Wang Jietao at the Wuhan Center, China Geological Survey provided eldwork and
logistical support and developed part of gure 1. We thank International Research Cent-
er on Karst in Guilin for additional logistical support. Mr. Zhang Yuanhai provided us
with information regarding regional caves while Dr. Qiang Li provided cave coordinates
for some of the caves examined in this study. Dr. Wu Zhuoting oered additional assis-
tance with some of the Mandarin names. Dr. Sergei Golovatch provided us with invalu-
able peer-review comments leading to the improvement of this manuscript. Fieldwork
was recognized as an Explorers Club ag expedition. e lead author was sponsored by
the National Natural Science Foundation of China (Grant no. 31801956 and Grant no.
41871039) for work related to species’ descriptions and manuscript preparation.
References
Barr Jr TC (1967) Observations on the ecology of caves. American Naturalist 101: 475–491.
https://doi.org/10.1086/282512
Barr Jr TC (1985) Pattern and process in speciation of trechine beetles in eastern North Amer-
ica (Coleoptera: Carabidae: Trechinae). Phylogeny and zoogeography of beetles and ants,
Junk, Dordrecht [1985 Sep 30]: 350–407.
Benedict EM (1979) A new species of Apochthonius Chamberlin from Oregon (Pseudoscorpio-
nida, Chthoniidae). Journal of Arachnology 7: 79–83.
Bernard EC, Soto-Adames FN, Wynne JJ (2015) Collembola of Rapa Nui (Easter Island) with
descriptions of ve endemic cave-restricted species. Zootaxa 3949: 239–267. https://doi.
org/10.11646/zootaxa.3949.2.6
Cao J, Yang D, Zhang C, Jiang Z (2007) Karst ecosystem of Guangxi Zhuang Autonomous
Region Constrained by Geological Setting: Relationship between carbonate rock exposure
and vegetation coverage. International Conference on Karst Hydrogeology and Ecosys-
tems, USA: 211–218.
Chen Z, Decu V, Juberthie C, Ueno SI (2001) China. In: Juberthie C, Decu V (Eds) Ency-
clopaedia Biospeologica, Part III, Société Internationale de Biospéologie, Moulis, France,
1763–1781.
Chevaldonné P, Lejeune C (2003) Regional warming-induced species shift in northwest Medi-
terranean marine caves. Ecology Letters 6: 371–379. https://doi.org/10.1046/j.1461-
0248.2003.00439.x
Christman MC, Culver DC, Madden MK, White D (2005) Patterns of endemism of the east-
ern North American cave fauna. Journal of Biogeography 32: 1442–1452. https://doi.
org/10.1111/j.1365-2699.2005.01263.x
Clarke AK (2006) Guangxi 2005 cave fauna report (China Caves Project). YRC Bulletin, Au-
tumn 2006.
Clements R, Sodhi NS, Schilthuizen M, Ng PK (2006) Limestone karsts of Southeast Asia:
imperiled arks of biodiversity. Bioscience 56: 733–742. https://doi.org/10.1641/0006-
3568(2006)56[733:LKOSAI]2.0.CO;2
Cave-dwelling millipedes, Guangxi, China 89
Culver DC, Master LL, Christman MC, Hobbs HH, 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
Deharveng L, Bedos A (2000) e cave fauna of Southeast Asia: origin, evolution and ecology.
In: Wilkens H, Culver DC, Humphreys WF (Eds) Ecosystems of the World 30: Subter-
ranean Ecosystems, Elsevier, Amsterdam, 603–632.
Deharveng L, Bréhier F, Bedos A, Tian MY, Li YB, Zhang F, Qin WG, Tan XF (2008) Mulun
and surrounding karsts (Guangxi) host the richest cave fauna of China. Subterranean Biol-
ogy 6: 75–79.
Elliott WR (1992). Fire ants invade Texas caves. American Caves Winter 5: 13.
Engho H, Golovatch SI, Short M, Stoev P, Wesener T (2015) Diplopoda – taxonomic over-
view. In: Minelli A (Ed) Treatise on Zoology-Anatomy, Taxonomy, Biology. e Myri-
apoda 2: 365–457.
Faille A, Tänzler R, Toussaint EF (2015) On the way to speciation: shedding light on the karstic
phylogeography of the microendemic cave beetle Aphaenops cerberus in the Pyrenees.
Journal of Heredity 106: 692–699. https://doi.org/10.1093/jhered/esv078
Ferreira RL, Horta LCS (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, Wynne JJ, Zhang F (2018) Two new species of cave-adapted pseudoscorpions (Pseu-
doscorpiones, Neobisiidae, Chthoniidae) from Guangxi, China. Journal of Arachnology
46: 345–354. https://doi.org/10.1636/JoA-S-17-063.1
Golovatch SI, Georoy JJ (2006) Review of the Southeast Asian millipede genus Pacidesmus
Golovatch, with the description of a new troglobitic species from southern China (Diplop-
oda: Polydesmida: Polydesmidae). Zootaxa 1325: 363–368. https://doi.org/10.11646/
zootaxa.1325.1.24
Golovatch SI, Georoy JJ, Mauriès JP (2006) Review of the millipede genus Hyleoglomeris Ver-
hoe, 1910 (Diplopoda, Glomerida, Glomeridae), with descriptions of new species from
caves in southeast Asia. Zoosystema 28: 887–915.
Golovatch SI, Georoy JJ, Mauriès JP, VandenSpiegel D (2007) Review of the millipede genus
Glyphiulus Gervais, 1847, with descriptions of new species from Aoutheast Asia (Diplop-
oda, Spirostreptida, Cambalopsidae). Part 1: the granulatus-group. Zoosystema 29: 7–49.
Golovatch SI, Georoy JJ, Mauriès JP, VandenSpiegel D (2009a) Review of the millipede genus
Eutrichodesmus Silvestri, 1910 (Diplopoda, Polydesmida, Haplodesmidae), with descrip-
tions of new species. ZooKeys 12: 1–46. https://doi.org/10.3897/zookeys.12.167
Golovatch, SI, Georoy, JJ, Mauriès JP, VandenSpiegel D (2009b) Review of the millipede fam-
ily Haplodesmidae, with descriptions of some new or poorly-known species (Diplopoda,
Polydesmida). ZooKeys 7: 1–53. https://doi.org/10.3897/zookeys.7.117
Golovatch SI, Georoy JJ, Mauriès JP (2010a) Two new species of the millipede genus Desmox-
ytes Chamberlin, 1923 (Diplopoda: Polydesmida: Paradoxosomatidae) from caves in
southern China. Arthropoda Selecta 19: 57–61. https://doi.org/10.15298/arthsel.19.2.01
Golovatch SI, Georoy JJ, Mauriès JP (2010b) Review of the millipede genus Pacidesmus Golo-
vatch, 1991, with descriptions of three new species from caves in southern China (Di-
plopoda: Polydesmida: Polydesmidae). Tropical Natural History 10: 159–169.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
90
Golovatch SI, Georoy JJ, Mauriès JP, VandenSpiegel D (2011a) Two new species of the mil-
lipede genus Hypocambala Silvestri, 1895 from China and Vietnam (Diplopoda: Spiro-
streptida: Cambalopsidae). Arthropoda Selecta 20: 167–174. https://doi.org/10.15298/
arthsel.20.3.03
Golovatch SI, Georoy JJ, Mauriès JP (2011b) New species of the millipede genus Glyphiulus
Gervais, 1847 from thee granulatus-group (Diplopoda, Spirostreptida, Cambalopsidae).
Arthropoda Selecta 20: 65–114. https://doi.org/10.15298/arthsel.20.2.01
Golovatch SI, Georoy JJ, Mauriès JP, VandenSpiegel D (2011c) New species of the mil-
lipede genus Glyphiulus Gervais, 1847 from the javanicus-group (Diplopoda: Spiros-
treptida: Cambalopsidae). Arthropoda Selecta 20: 149–165. https://doi.org/10.15298/
arthsel.20.3.02
Golovatch SI, Liu WX, Georoy JJ (2012a) Review of the millipede genus Hyleoglomeris Verho-
e, 1910 in China, with descriptions of new species (Diplopoda, Glomerida, Glomeridae).
Zootaxa 3358: 1–27. https://doi.org/10.11646/zootaxa.3358.1.1
Golovatch SI, Li Y, Liu W, Georoy JJ (2012b) ree new cavernicolous species of dragon mil-
lipedes, genus Desmoxytes Chamberlin, 1923, from southern China, with notes on a formal
congener from the Philippines (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys
185: 1–18. https://doi.org/10.3897/zookeys.185.3082
Golovatch SI, Li Y, Liu W, Georoy JJ (2012c) One new and two little-known species of the
millipede family Polydemsidae from southern China (Diplopoda: Polydesmda). Arthropo-
da Selecta 21: 131–136. https://doi.org/10.15298/arthsel.21.2.02
Golovatch SI, Georoy JJ (2014) On some new or poorly-known species of the millipede fam-
ily Polydesmidae from southern China (Diplopoda: Polydesmida). Russian Entomological
Journal 23: 91–105. https://doi.org/10.15298/rusentj.23.2.01
Golovatch SI (2015) Cave Diplopoda of southern China with reference to millipede diversity
in Southeast Asia. ZooKeys 510: 79–94. https://doi.org/10.3897/zookeys.510.8640
Golovatch SI, Georoy JJ, Mauriès JP, VandenSpiegel D (2015) Review of the millipede genus
Eutrichodesmus Silvestri, 1910, in China, with descriptions of new cavernicolous species
(Diplopoda, Polydesmida, Haplodesmidae). Zookeys 505: 1–34. https://doi.org/10.3897/
zookeys.505.9862
Harley GL, Polk JS, North LA, Reeder PP (2011) Application of a cave inventory system to
stimulate development of management strategies: e case of west-central Florida, USA.
Journal of Environmental Management 92: 2547–2557. https://doi.org/10.1016/j.jenv-
man.2011.05.020
Harvey MS, Wynne JJ (2014) Troglomorphic pseudoscorpions (Arachnida: Pseudoscorpiones)
of northern Arizona, with descriptions of two new short-range endemic species. Journal of
Arachnology 42: 205–219. https://doi.org/10.1636/K14-34.1
Howarth FG (1980) e Zoogeography of Specialized Cave Animals: A Bioclimatic Model.
Evolution 34: 394–406. https://doi.org/10.1111/j.1558-5646.1980.tb04827.x
Howarth FG (1982) Bioclimatic and geological factors governing the evolution and distribu-
tion of Hawaiian cave insects. Entomologia Generalis 8: 17–26.
Howarth FG (1983) Ecology of cave arthropods. Annual Review of Entomology 28: 365–389.
https://doi.org/10.1146/annurev.en.28.010183.002053
Cave-dwelling millipedes, Guangxi, China 91
Howarth FG, James SA, McDowell W, Preston DJ, Imada CT (2007). Identication of roots
in lava tube caves using molecular techniques: Implications for conservation of cave ar-
thropod faunas. Journal of Insect Conservation 11: 251–261. https://doi.org/10.1007/
s10841-006-9040-y
Jiang X, Guo X, Chen H, Xie Z (2018) Four new species of the Glyphiulus javanicus group from
southern China (Diplopoda, Spirostreptida, Cambalopsidae). ZooKeys 741: 155–179.
https://doi.org/10.3897/zookeys.741.23223
Katz AD, Taylor SJ, Davis MA (2018) At the conuence of vicariance and dispersal: Phyloge-
ography of cavernicolous springtails (Collembola: Arrhopalitidae, Tomoceridae) codistrib-
uted across a geologically complex karst landscape in Illinois and Missouri. Ecology and
Evolution 8: 10306–10325. https://doi.org/10.1002/ece3.4507
Lindsey AA (1951) Vegetation and habitats in a southwestern volcanic area. Ecological Mono-
graphs 21: 227–253. https://doi.org/10.2307/1943559
Likhitrakarn N, Golovatch SI, Srisonchai R, Panha S (2017) A new species of Trichopeltis Po-
cock, 1894 from southern China, with a checklist and a distribution map of Trichopeltis
species (Diplopoda, Polydesmida, Cryptodesmidae). ZooKeys 725: 123–137. https://doi.
org/10.3897/zookeys.725.22014
Liu WX, Tian MY (2013) Four new cavernicolous species of the millipede genus Eutrichodes-
mus Silvestri, 1910 from southern China (Diplopoda: Polydesmida: Haplodesmidae).
Zootaxa 3734: 281–291. https://doi.org/10.11646/zootaxa.3734.2.11
Liu WX, Golovatch SI, Tian MY (2014) A review of the dragon millipede genus Desmoxytes
Chamberlin, 1923 in China, with descriptions of four new species (Diplopoda: Polydesmi-
da: Paradoxosomatidae). ZooKeys 448: 9–26. https://doi.org/10.3897/zookeys.448.8081
Liu WX, Tian MY (2015a) A checklist of millipede genus Hyleoglomeris Verhoe, 1910 in
mainland China, with descriptions of seven new species (Diplopoda, Glomerida, Glomeri-
dae). Zootaxa 4032: 103–116. https://doi.org/10.11646/zootaxa.4032.1.5
Liu WX, Tian MY (2015b) Occurrence of the millipede genus Piccola Attems, 1953 in Chi-
na(Diplopoda: Polydesmida: Paradoxosomatidae). Zootaxa 3904: 403–408. https://doi.
org/10.11646/zootaxa.3904.3.5
Liu W X, Tian MY (2015c) Two new cave-dwelling species of the millipede genus Paracortina
Wang & Zhang, 1993 from southern China (Diplopoda, Callipodida, Paracortinidae).
ZooKeys 517: 123–140. https://doi.org/10.3897/zookeys.517.9949
Liu WX, Golovatch SI, Tian MY (2016) Six new species of dragon millipedes, genus Desmoxy-
tes Chamberlin, 1923, mostly from caves in China (Diplopoda, Polydesmida, Paradoxoso-
matidae). ZooKeys 557: 1–24. https://doi.org/10.3897/zookeys.577.7825
Liu WX, Golovatch SI, Wesener T, Tian MY (2017a) Convergent Evolution of Unique Mor-
phological Adaptations to a Subterranean Environment in Cave Millipedes (Diplopoda).
PLoS One 12: e0170717. https://doi.org/10.1371/journal.pone.0170717
Liu WX, Golovatch SI, Wesener T (2017b) Four new species of the millipede genus Eutrichodes-
mus Silvestri, 1910 from Laos, including two with reduced ozopores (Diplopoda, Polydes-
mida, Haplodesmidae). ZooKeys 660: 43–65. https://doi.org/10.3897/zookeys.660.11780
Loksa I (1960) Einige neue Diplopoden- und Chilopodenaren aus chinesischen Höhlen. Acta
Zoologica Academiae Scientiarum Hungaricae 6: 135–148.
Weixin Liu & J. Judson Wynne / Subterranean Biology 30: 57–94 (2019)
92
Mauriès JP, Nguyen Duy-Jacquemin M (1997) Nouveaux craspedosomides et glyphiulides cav-
ernicoles de Chine (Diplopoda). Mémoires de Biospéologie 24: 49–62.
Mammola S, Goodacre SL, Isaia M (2018) Climate change may drive cave spiders to extinc-
tion. Ecography 41: 233–243. https://doi.org/10.1111/ecog.02902
Mesibov R, Taylor RJ, Brereton RN (1995) Relative eciency of pitfall trapping and hand-col-
lecting from plots for sampling of millipedes. Biodiversity and Conservation 4: 429–439.
https://doi.org/10.1007/BF00058426
Minelli A (2015) Treatise on Zoology-Anatomy, Taxonomy, Biology. e Myriapoda 2. Brill.,
482 pp. https://doi.org/10.1163/9789004188273
Mockford EL, Wynne JJ (2013) Genus Cyptophania Banks (Psocodea: Lepidopsocidae): Unique
features, augmented description of the generotype, and descriptions of three new species.
Zootaxa 3702: 437–449. https://doi.org/10.11646/zootaxa.3702.5.3
Monro AK, Bystriakova N, Fu L, Wen F, Wei Y, (2018) Discovery of a diverse cave ora in
China. PloS One 13: e0190801. https://doi.org/10.1371/journal.pone.0190801
Nitzu E, Vlaicu M, Giurginca A, Meleg IN, Popa I, Nae A, Baba Ş (2018) Assessing preservation
priorities of caves and karst areas using the frequency of endemic cave-dwelling species. Inter-
national Journal of Speleology 47: 43–52. https://doi.org/10.5038/1827-806X.47.1.2147
Northup DE, Welbourn WC (1997) Life in the twilight zone – Lava tube ecology, natural
history of El Malpais National Monument. New Mexico Bureau of Mines and Mineral
Resources Bulletin 156: 69–82.
Peck SB (1970) e terrestrial arthropod fauna of Florida caves. Florida Entomologist 53:
203–207. https://doi.org/10.2307/3493189
Price L (2016) An introduction to some cave fauna of Malaysia and ailand. Acta Carsologica
33: 311–317. https://doi.org/10.2307/3493189
Ravbar N (2016) e earliest Chinese karstologist Xu Xiake. Acta Carsologica 32: 243–254.
https://doi.org/10.3986/ac.v32i1.376
Reddell JR (1994) e cave fauna of Texas with special reference to the western Edwards Pla-
teau. In: Elliott WR, Veni G (Eds) e caves and karst of Texas. National Speleological
Society, Huntsville, 31–50.
Reeves WK (1999) Exotic species of North American Caves. In: Henderson K (Ed.) Proceed-
ings of the 1999 National Cave and Karst management symposium, Chattanooga, Tennes-
see: National Cave & Karst Management Symposia, 164–166.
Sket B (2008) Can we agree on an ecological classication of subterranean animals? Journal of
Natural History 42: 1549–1563. https://doi.org/10.1080/00222930801995762
Srisonchai R, Engho H, Likhitrakarn N, Panha S (2018) A revision of dragon millipedes I:
genus Hylomus Chamberlin, 1923, with the description of eight new species (Diplopoda,
Polydesmida, Paradoxosomatidae). ZooKeys 761: 1–177. https://doi.org/10.3897/zook-
eys.761.24214
Stone FD, Howarth FG (2007) Hawaiian cave biology: status of conservation and manage-
ment. In: Proceedings of the 2005 National Cave and Karst Management Symposium;
Albany, New York [October 31–November 4, 2005], 21–26.
Sweeting MM (1995) Karst in China. Its geomorphology and environment. Springer Verlag,
New York, 265 pp. https://doi.org/10.1007/978-3-642-79520-6
Cave-dwelling millipedes, Guangxi, China 93
Taiti S, Wynne JJ (2015) e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Is-
land), with descriptions of two new species. ZooKeys 515: 27–49. https://doi.org/10.3897/
zookeys.515.9477
Tanalgo KC, Tabora JA, Hughes AC (2018) Bat cave vulnerability index (BCVI): A holistic
rapid assessment tool to identify priorities for eective cave conservation in the tropics.
Ecological Indicators 89: 852–860. https://doi.org/10.1016/j.ecolind.2017.11.064
Taylor SJ, Krejca J, Smith JE, Block VR, Hutto F (2003) Investigation of the potential for
red imported re ant (Solenopsis invicta) impacts on rare karst invertebrates at Fort Hood,
Texas: A eld study. In: Illinois Bexar County Karst Invertebrates Draft Recovery Plan
Natural History Survey. Center for Biodiversity Technical Report, vol. 28, 1–153.
Tian MY (2011) A new subgenus and two new species of the troglobitic genus Dongodytes
Deuve from Guangxi, China (Coleoptera, Carabidae). Subterranean Biology 8: 57–64.
https://doi.org/10.3897/subtbiol.8.1232
Trajano E (2000) Cave faunas in the Atlantic tropical rain forest: Composition, ecology and con-
servation. 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. https://doi.org/10.1111/j.1365-
2664.2009.01630.x
Wynne JJ (2013) Inventory, conservation and management of lava tube caves at El Malpais
National Monument, New Mexico. Park Science 30: 45–55.
Wynne JJ, Shear WA (2016) A new millipede (Austrotyla awishashola, n. sp., Diplopoda, Chor-
deumatida, Conotylidae) from New Mexico, USA, and the importance of cave moss gardens
as refugial habitats. Zootaxa 4084: 285–292. https://doi.org/10.11646/zootaxa.4084.2.8
Wynne JJ, Bernard EC, Howarth FG, Sommer S, Soto-Adames FN, Taiti S, Mockford EL,
Horrocks M, Pakarati L, Pakarati-Hotus V (2014) Disturbance relicts in a rapidly chang-
ing World: e Rapa Nui (Easter Island) factor. BioScience 64: 711–718. https://doi.
org/10.1093/biosci/biu090
Wynne JJ, Sommer S, Howarth FG, Dickson BG, Voyles KD (2018) Capturing arthropod
diversity in complex cave systems. Diversity and Distributions 24: 1478–1491. https://
doi.org/10.1111/ddi.12772
Wynne JJ, Howarth FG, Sommer S, Dickson BG (2019) Fifty years of cave arthropod sam-
pling: techniques and best practices. International Journal of Speleology 48: 33–48.
https://doi.org/10.5038/1827-806X.48.1.2231
Zhang CZ, Li ZY (1981) Bilinguidae family nov. of Diplopoda in karst landform of south
China. Acta Zootaxonomica Sinica 6: 373–377. [In Chinese]
Zhao Y, Gluesenkamp A, Fenolio D, Soares D, Niemiller M, Bichuette M-E, Chakrabarty P
(in press) e species diversity and conservation of caveshes in China. In: Wynne JJ (Ed.)
Cave Ecology: Drivers of Diversity and Speciation. NOVA Science Publishers, NY.
Available via license: CC BY 4.0
Content may be subject to copyright.
