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259
The millipede family Polydesmidae Leach, 1816 (Diplopoda,
Polydesmida) from Vietnam, with a description of a new
cavernicolous species
Anh D. Nguyen1,2 , Tam T. T. Vu1, Katsuyuki Eguchi3,4
1 Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18, Hoangquocviet Rd., Caugiay District, Hanoi, Vietnam
2 Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18, Hoangquocviet Rd., Caugiay District, Hanoi, Vietnam
3 Graduate School of Science, Tokyo Metropolitan University, Minami-osawa 1-1, Hachioji, Tokyo 192-0397, Japan
4 Department of International Health and Medical Anthropology, Institute of Tropical Medicine, 1-12-4 Sakamoto, Nagasaki University, Nagasaki, 852-8523, Japan
Corresponding author: Anh D. Nguyen (ducanh410@yahoo.com, ndanh@iebr.vast.vn)
Copyright: © Anh D. Nguyen et al.
This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International –
CC BY 4.0).
Research Article
Abstract
The millipede family Polydesmidae Leach, 1816 is reviewed in the scope of the Vietnam-
ese fauna. The distribution of the species, Polydesmus vietnamicus Nguyen, 2009 is ex-
tended northward to Ha Giang Province. A new cavernicolous polydesmid, Pacidesmus
tuachua sp. nov., is described from two caves in northwestern Vietnam, representing the
provided for both species, a revised key is presented to all 12 species of Pacidesmus
Golovatch, 1991, as well as a key to all eight genera of Asian Polydesmidae.
Key words: Asia, cave fauna, COI sequence, diversity, new species, taxonomy
Introduction
The millipede family Polydesmidae Leach, 1816 is almost strictly Holarctic,
consisting of more than 60 nominal genera or subgenera and nearly 400 spe-
cies and subspecies (Hoffman 1980; Golovatch 1991). The family is mostly
distributed in the Mediterranean area, whereas Central and East Asia, as well as
the entire Nearctic Region, show lower generic and, to a lesser degree, species
diversity (Golovatch 1991; Djursvoll et al. 2001). Only a few macropolydesmid
genera are found in Asia and Indochina including Epanerchodus Attems, 1901,
Pacidesmus Golovatch, 1991, Polydesmus Latreille, 1802–1803, Nipponesmus
Chamberlin & Wang, 1953, Gleninea Turk, 1945, Jaxartes Verhoeff, 1930, Schizo-
turanius Verhoeff, 1931, and Uniramidesmus Golovatch & Mikhaljova, 1979
(Golovatch 1991; Geoffroy and Golovatch 2004; Mikhaljova 2004; Golovatch
Liu and Golovatch 2020). In addition, a fossil species of the rather small west-
ern to central European genus Propolydesmus Verhoeff, 1895 has recently been
described from the mid-Cretaceous amber of Myanmar (burmite, 99–100 Mya)
(Su et al. 2023).
Academic editor: Dragan Antić
Received:
31 October 2023
Accepted:
11 January 2024
Published:
30 January 2024
ZooBank: https://zoobank.org/
A5D5D928-CE84-4E13-BC31-
ED2393B3403D
Citation: Nguyen AD, Vu TTT,
Eguchi K (2024) The millipede
family Polydesmidae Leach, 1816
(Diplopoda, Polydesmida) from
Vietnam, with a description of a new
cavernicolous species. ZooKeys 1190:
259–280. https://doi.org/10.3897/
zookeys.1190.114958
ZooKeys 1190: 259–280 (2024)
DOI: 10.3897/zookeys.1190.114958
260
ZooKeys 1190: 259–280 (2024), DOI: 10.3897/zookeys.1190.114958
Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
Vietnam has been known to harbour a rich fauna of millipedes with about
250 recorded species (Enghoff et al. 2004 and updated). Many new species
have been discovered recently, including cave millipedes, e.g. Hyleoglomer-
is alba Nguyen et al., 2022 (Kuroda et al. 2022), Tylopus nguyeni Golovatch,
2019, and Hylomus srisonchai Golovatch, 2019 (Golovatch 2019), but only one
polydesmid has hitherto been revealed: Polydesmus vietnamicus Nguyen, 2009
from Tam Dao National Park, Vinh Phuc Province (Nguyen 2009).
The present paper updates the knowledge of the family Polydesmidae in Viet-
nam, with the description of a new cavernicolous species found in northwest-
ern Vietnam. An updated key to all 12 Pacidesmus species is also presented,
as well as a key to all eight genera of Polydesmidae reported so far from Asia.
Materials and methods
Millipede specimens were hand-collected from forests and caves in northern
Vietnam and preserved in 85–90% ethanol. Morphological characters were
investigated with an Olympus SZX16 stereomicroscope. Gonopods were dis-
sected for morphological examination and photographed. Colour images were
taken at various focal planes using a Nikon imaging system (Nikon-Br) coupled
with a SMZ800N Nikon stereomicroscope. UV images were taken using a Sony
a6000 digital camera attached to the aforementioned SMZ800N Nikon stereo-
Helicon Focus version 7.0 and assembled in Adobe Photoshop CS6. Scanning
electron microscope (SEM) images were taken using the system Prisma E
Total DNA was extracted using Qiagen DNeasy Blood and Tissue Kits. A
680-bp fragment of the mitochondrial gene, cytochrome c oxidase subunit I
(COI
and HCO2198 (Folmer et al. 1994). Polymerase chain reaction (PCR) condi-
GenBank accession numbers.
Morphological terminology follows Liu and Golovatch (2020). All specimens
reported here, including types, are deposited in the Institute of Ecology and
Biological Resources (IEBR), Vietnam Academy of Science and Technology,
Hanoi, Vietnam.
Abbreviation
IEBR-Myr Institute of Ecology and Biological Resources, Myriapod collection.
Results
Taxonomy
Order Polydesmida Pocock, 1887
Family Polydesmidae Leach, 1816
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
Genus Polydesmus Latreille, 1802–1803
Type species. Julus complanatus Linnaeus, 1761, by monotypy.
Remarks. Polydesmus is certainly the largest genus within the family
Polydesmidae, with over 200 species and subspecies which mainly occur in
Europe and the Mediterranean, west of the central Caucasus (Hoffman 1980;
Djursvoll et al. 2001). A few species have been found in the Oriental Region, these
being Polydesmus japonicus Miyosi, 1956, P. miyosii Murakami, 1966, P. tanakai
Murakami, 1970, and P. tangonis Murakami, 1973 from Japan; P. moorei Poco-
ck, 1895 and P. paludicola Pocock, 1895 from eastern China; P. liber Golovatch,
1991 from Hong Kong, southern China (Golovatch 1991); and P. vietnamicus
Nguyen, 2009, the only species of the genus from northern Vietnam (Nguyen
2009). The two old species of Pocock (1895) from mainland China are only
provisionally to be assigned to Polydesmus, as both require revision. The distri-
bution of Polydesmus seems to be amphi-Palaearctic (Golovatch 1991).
Polydesmus vietnamicus Nguyen, 2009
Figs 1, 2–8
Materials examined. – Vinh Phuc Province • 1 ♂, 1 ♀; Vinh Phuc Prov-
ince, Tam Dao National Park, near town; 1,000 m a.s.l.; 1 March 2005; Anh D.
Nguyen leg.; natural secondary forest • 1 ♂; Vinh Phuc Province, Tam Dao Na-
tional Park, on the way to Thac Bac waterfall; 1,000 m a.s.l.; 22 March 2005;
Anh D. Nguyen leg.; bamboo forest, near stream • 1 ♂, 3 ♀s, 1 juvenile; Vinh
Phuc Province, Tam Dao National Park, around the town; 900–1,000 m a.s.l.;
15–18 October 2010; Anh D. Nguyen leg.; mixed forest; IEBR-Myr 967 • 3 ♀♀;
Tam Dao National Park, on way to Tam Dao 2; 1,100 m a.s.l.; 25 February 2017;
Anh D. Nguyen leg.; natural forest; IEBR-Myr 604 – Ha Giang Province • 2 ♂♂,
1 ♀; Bac Me Natural Reserve, Lac Nong commune, Ban Khen; 22°45'30.8"N,
105°14'04.5"E; 11 December 2019; Anh D. Nguyen leg.; regenerated forest; IE-
BR-Myr 808.
Diagnosis. A typical polydesmid with 20 body rings and three transverse
rows of bosses with setae on metaterga. Gonopodal solenomere rather well
developed, conspicuously shaped. Endomere elongate and strongly falcate, di-
rected caudally, starting laterally and basally of recurvature point of seminal
groove, set off from femorite by a sulcus, with a pair of strong teeth at about
midway (mt). Seminal groove largely mesal, crossing the femorite diagonally,
terminal lateral loop relatively short and turning around a distofemoral process
(ap). Solenomere (sl
The species differs from the morphologically particularly similar Polydesmus
liber Golovatch, 1991 in being larger (33.0–38.4 mm vs 21.0–23.0 mm in length)
and in the gonopod endomere (with a pair of teeth at about its midlength vs
with two pairs of moderate teeth at 1/3 and 2/3 of its length).
It is particularly noteworthy that all East and Southeast Asian species un-
doubtedly belonging to Polydesmus, however few, share the symplesiomorphy
of densely setose gonopod coxites, which contrasts with very poorly setose
ones observed in the much more numerous western Palaearctic counterparts
(Golovatch 1991).
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
DNA barcode. The COI fragment (660 bp) was uploaded to GenBank with
accession numbers PP118038 and PP118039. Polydesmus vietnamicus
has a close COI identity to Pseudopolydesmus pinetorum (Bollman, 1888)
(MT739870) and Pseudopolydesmus serratus (Say, 1821) (MT739862), with
89.8% (query coverage 83%) and 88.71% (query coverage 83%), respectively.
Remarks. This species was previously known from only its type locality, Tam
Dao National Park (Nguyen 2009). Currently, its distribution is extended north-
between the type specimens and those samples collected in Ha Giang.
Genus Pacidesmus Golovatch, 1991
Type species. Pacidesmus shelleyi Golovatch, 1991, by original designation.
Remarks. Pacidesmus contains 12 species found in southern China and north-
ern Thailand (listed below). While the type species, P. shelleyi Golovatch, 1991,
is known from forest litter at 2,200 m a.s.l. on Mount Doi Inthanon in northern
Thailand (Golovatch 1991), the remaining species seem to be troglobionts re-
stricted to caves in southern China, especially Guangxi and Guizhou provinces
(Golovatch et al. 2010; Golovatch and Geoffroy 2014; Liu and Golovatch 2020).
Figure 1. Records of polydesmid species in Vietnam and of all known Pacidesmus species.
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
Pacidesmus tuachua sp. nov.
https://zoobank.org/CB8E9D35-3365-4416-81CA-589284A6D42F
Figs 1, 9–14
Materials examined. Holotype. • ♂; Dien Bien Province, Tua Chua Dis-
trict, Xa Nhe commune, Xa Nhe cave; 600 m a.s.l.; 21°52'37"N, 103°24'48"E; 12
April 2022; Anh D. Nguyen leg.; IEBR-Myr 951H.
Paratypes. – Dien Bien Province • 5 ♂♂, 10 ♀♀; Tua Chua District, Xa
Nhe commune, Kho Chua La cave; 600 m a.s.l.; 21°52'36.9"N, 103°24'47.9"E; 12
January 2021; Anh D. Nguyen leg.; IEBR-Myr 899 • 3 ♂♂, 4 ♀♀; Tua Chua District,
Xa Nhe commune, Kho Chua La cave; 600 m a.s.l.; 21°52'36.9"N, 103°24'47.9"E;
12 January 2021; Anh D. Nguyen leg.; IEBR-Myr 900 • 4 ♂♂, 5 ♀♀; Tua Chua
District, Xa Nhe commune, Kho Chua La cave; 600 m a.s.l.; 21°52'36.9"N,
103°24'47.9"E; 12 April 2022; Anh D. Nguyen leg.; IEBR-Myr 951P • 4 ♂♂, 5 ♀♀;
Tua Chua District, Xa Nhe commune, Xa Nhe cave; 600 m a.s.l.; 21°52'37"N,
103°24'48"E; 12 April 2022; Anh D. Nguyen leg.; IEBR-Myr 952 • 2 ♂♂, 4 ♀♀; Tua
Chua District, Xa Nhe commune, Kho Chua La cave; 600 m a.s.l.; 21°52'36.9"N,
103°24'47.9"E; 12 April 2022; Anh D. Nguyen leg.; IEBR-Myr 953.
Figure 2. Polydesmus vietnamicus Nguyen, 2009 from Ha Giang Province (IEBR-Myr 808) A–C ♂. Anterior part of body
A dorsal view B lateral view C ventral view D ♀, anterior part of body, ventral view. Scale bars: 1 mm.
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Diagnosis. The new species can be distinguished from its congeners by a
combination of the following features: unpigmented colouration, small size
(midbody width <4.0 mm), head narrower than collum, absence of sphaero-
trichomes, lateral budges on male prefemora, subfalcate gonopod telopo-
dite, absence of exomere, endomere with an acute triangular process distally
and a broad triangular process medially, and endomere tip slightly and un-
The species is truly cavernicolous, characterized by white or unpigment-
ed colour and living within a cave. As a troglobiont species, it groups with
all 12 troglobiont or troglophile congeners from China (Table 1). However,
this species differs from all of these, except P. bifidus from the Hengli Xin
Don Cave, Guangxi Province in the absence of an exomere and the gonopod
telopodite showing no additional processes; the endomere also has two ad-
ditional processes, and the tip of the endomere bears two tiny teeth. The
new species is similar to P. bifidus in having a troglomorphic appearance,
the absence of an exomere, and a bifid tip of the endomere, but it differs in
having two tiny teeth at the tip of the endomere tip (vs two long processes
in P. bifidus).
Figure 3. Polydesmus vietnamicus Nguyen, 2009 from Ha Giang Province (IEBR-Myr 808) ♂ A midbody segment 8–11,
dorsal view B posterior part of body, dorsal view C posterior part of body, lateral view D posterior part of body, ventral
view. Scale bars: 1 mm.
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
Figure 4. Polydesmus vietnamicus Nguyen, 2009 from Ha Giang Province (IEBR-Myr 808) ♂ and ♀ under the UV light
A ♂, anterior part of body, dorsal view B ♂, anterior part of body, ventral view C ♀, anterior part of body, ventral view D ♂,
midbody segment 8-10, dorsal view. Scale bars: 1 mm.
Table 1. List of all known species of the genus Pacidesmus Golovatch, 1991.
Species Localities
Pacidesmus armatus Golovatch, Geoffroy & Mauriès, 2010 China, Guangxi Prov., Huanjiang, Cave Xiao Lan Dong (Golovatch et al. 2010)
Pacidesmus bedosae Golovatch, Geoffroy & Mauriès, 2010 China, Guangxi Prov., Huanjiang, Cave Dong Tu Dong (Golovatch et al. 2010)
Pacidesmus bidus Golovatch & Geoffroy, 2014 China, Guangxi Prov., Cave Hengli Xin Dong near Fengshan (Fengshan Xian)
(Golovatch and Geoffroy 2014)
Pacidesmus martensi Golovatch & Geoffroy, 2006 China, Guizhou Prov., Qianxi County, Hong Lin Town, Ishui Luo Dong Cave China, Guizhou Prov.,
Dafang County, Yangzhamba Village, Hei Dong Cave China, Guizhou Province, Qianxi County,
Honglin Town, Jisha Village, I Dong Cave (Golovatch et al. 2007; Golovatch and Geoffroy 2006)
Pacidesmus shelleyi Goolovatch, 1991 Thailand, Chieng Mai Province, Doi Inthanon National Park (Golovatch 1991)
Pacidesmus sinensis (Golovatch & Hoffman, 1989) China, Guizhou Province, Ziyun County, Getuhe National Geopark, Suidao Dong Cave (Liu and
Golovatch 2020) A cave in Guizhou Province and Cave Kaikou Dong, Zhenning County, Guizhou
Province, China (Loksa 1960; Golovatch and Hoffman 1989; Chen and Meng 1990)
Pacidesmus superdraco Golovatch, Geoffroy & Mauriès, 2006 Cave Laitai Dong, Libo County, Guizhou Province (Golovatch et al. 2007)
Pacidesmus tiani Golovatch, Geoffroy & Mauriès, 2010 China, Guangxi Prov., Huanjiang, Cave Gang Lai Dong (Golovatch et al. 2010)
Pacidesmus tridus Golovatch & Geoffroy, 2014 China, Guangxi Prov., Guilin County, Grotte des Squelettes (Golovatch and Geoffroy 2014).
Pacidesmus trilobatus Liu & Golovatch, 2020 China, Yunnan Province, Maguan County, Pojiao Town, Dayan Dong Cave China, Yunnan Province,
Wenshan County, Liujing Town, Laozhai Village, I Dong Cave (Liu and Golovatch 2020).
Pacidesmus uncatus Liu & Golovatch, 2020 China, Yunnan Province, Qujing City, Zhanyi County, Tianshengqiao Dong Cave
(Liu and Golovatch 2020)
Pacidesmus whitteni Liu & Golovatch, 2020 China, Guangxi Zhuang Autonomous Region, Fengshan County, Jinya Town, Hangdong Village,
I Dong Cave (Liu and Golovatch 2020)
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The new species is assigned to Pacidesmus because of the following char-
acters: the seminal groove starts mesally, then recurves laterad at the base of a
particularly prominent endomere branch to enter an accessory seminal cham-
ber that opens on a setose pulvillus; endomere bears additional processes.
Etymology.
based on the “Tua Chua” district where the two caves are located.
Description. Holotype length ca 16.3 mm, width of midbody pro- and meta-
zonae 1.0 mm and 1.5 mm, respectively. In width, head < collum < segment 3 = 4
< 2 < 5 = 15, thereafter body gradually tapering towards telson (Figs 9, 10, 12). Co-
louration in alcohol rather uniformly white (Figs 8, 9). Body with 20 segments. An-
tennae long and only slightly clavate, possibly reaching past segment 3 if stretch-
ing laterally; antennomere 3 longest, approximately 1.3× longer than subequal
antennomeres 4–6; antennomeres 5 and 6 each with a small, compact, distodor-
sal group of bacilliform sensilla; antennomere 7 with a minute dorsoparabasal
cone and a distodorsal group of microscopic sensilla (Figs 9A–C, 11C).
Paraterga (Figs 9, 10, 12A–C) strongly developed, set high, starting with col-
lum, dorsum faintly convex; paraterga mostly weakly upturned above dorsum.
Caudolateral corner of paraterga acute, postcollum ones extending increas-
ingly past rear tergal margin, especially so in segments 16–18. All poreless
segments with three incisions, all pore-bearing ones with four minute incisions
at lateral margin. Front margins of metaterga narrowly bordered and forming
distinct shoulders.
Figure 5. Polydesmus vietnamicus Nguyen, 2009 from Ha Giang Province (IEBR-Myr 808) ♂, under the UV light A posterior
part of body, dorsal view B posterior part of body, ventral view C telson, dorsal view D telson, ventral view. Scale bars: 1 mm.
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Ozopores evident, dorsal, located in front of posteriormost marginal incision
of paraterga 5, 7, 9, 10, 12–13, 15–19.
Metatergal sculpture typical, poorly developed, obliterate, with three trans-
verse rows of typical (= polydesmid), setigerous, polygonal bosses. Tergal
setae short, slightly longer only on collum, simple, often obliterate. Stricture
between pro- and metazona wide, shallow and nearly smooth. Limbus exceed-
ingly thin, microdenticulate (Fig. 12A–C). Pleurosternal carinae absent.
Epiproct (Figs 11A, B, 12C, D) short, conical, pre-apical lateral papillae ev-
ident. Hypoproct (Figs 11A, 12D) subtriangular; distolateral setiferous knobs
small, but distinct and well separated.
Legs generally long and slender, apparently slightly incrassate, approximate-
ly 1.7–1.8× as long as midbody height, densely setose, almost all setae sim-
ple, poorly branching setae with minute, distal, side branchlets only on slender
prefemora, latter devoid of lateral bulges.
Gonopods (Figs 11D, 12, 13) characteristically subfalcate (vs suberect in all
other congeners). with large, rectangular coxites (co), with a few long setae
Figure 6. Polydesmus vietnamicus Nguyen, 2009 from Ha Giang Province (IEBR-Myr 808) ♂, segment 7 with gonopods
under the normal (A, B) and UV (C, D) light A, C posterior views B, D anterior views. Scale bars: 1 mm.
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ca) as usual. Telopodite elongate,
stout, strongly falcate or C-shaped; prefemorite (pref) densely setose; semi-
nal groove starting mesally, then recurving laterad to run to the opening on a
hairy pulvillus. Endomere (en) with two additional processes, a shorter, larger,
broader triangular process at midlength (p1), and a longer, acuter, triangular
process at ¾ length (p2-
er branch. Neither an exomere nor a clivus.
Figure 7. Polydesmus vietnamicus Nguyen, 2009 from Ha Giang Province (IEBR-Myr 808) ♂, right gonopod under the UV
light A ventral view B lateral view C dorsal view D mesal view. Scale bars: 1 mm.
A B
C D
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Figure 8. Polydesmus vietnamicus Nguyen, 2009 from Ha Giang Province (IEBR-Myr 808) ♂, right gonopod A mesal view
B lateral view. Abbreviations: co = coxite; pref = prefemorite; ca = cannula; en = endomere; sl = solenomere; mt = midway
teeth; ap = additional process. Scale bars: 0.5 mm.
Figure 9. Pacidesmus tuachua sp. nov., holotype ♂ (IEBR-Myr 951) A anterior part of body, dorsal view B anterior part of
body, lateral view C anterior part of body, ventral view D segments 8–10, dorsal view. Scale bars: 1 mm.
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Female. Slightly larger than male, length ca 16.8 mm, width of pro- and meta-
zona about 1.1 mm and 1.7 mm, respectively. Paraterga slightly less strongly
-
ly elevated. Epigynal ridge low.
DNA barcode. Two COI sequences (661 bp) were uploaded to the GenBank
with the accession numbers PP118040 and PP118041. The new species has
a close COI identity to Epanerchodus koreanus (NC051495) at 88.72% (query
coverage 97%).
Habitat. This species is to be considered a true troglobiont because it shows
the typical morphological features of a cave-dweller. It was collected exclusively
in the dark zone of the caves as described below. Kho Chua La and Xa Nhe caves
are both located close together, approximately 500 m in distance. These caves
are at the centre of the Xa Nhe commune, Dien Bien Province, northwestern Viet-
nam. The two caves are tunnel-like: they are high (15–20 m), wide (15–20 m),
pools. Several other millipede species have been found in these caves, including
Glyphiulus sp. (Spirostreptida, Glyphiulidae) and Eutrichodesmus sp. (Polydesmi-
da,: Haplodesmidae). The new species was found >1000 m from the entrance.
Kho Chua La and Xa Nhe caves are located on the Tua Chua karst plateau in
northeastern Dien Bien Province, northwestern Vietnam. The natural area is about
68,414 ha, and 70% of this area is composed of limestone mountains, which are
known for their layers of majestic rugged rock and unique natural landscape.
The karst region contains many stunning and well-known caves, such as Kho
Chua La, Tham Khem, Hau Chua, Xa Nhe, and Pe Rang Ki (Nguyen et al. 2022).
Furthermore, the Tua Chua karst plateau of northwestern Vietnam is close to the
Figure 10. Pacidesmus tuachua sp. nov., holotype ♂ (IEBR-Myr 951) A segments 8–10, ventral view B–D posterior part of
body, dorsal, vental and lateral views, respectively. Scale bars: 1 mm.
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Figure 11. Pacidesmus tuachua sp. nov., holotype ♂ (IEBR-Myr 951) A telson, ventral view B telson, lateral view C right
antenna, anterior view D gonopods in situ under UV light, ventral view. Scale bars: 1 mm.
Figure 12. Pacidesmus tuachua sp. nov., holotype ♂ (IEBR-Myr 951) A anterior part of body, under the UV light, dorsal view
B segments 8–10, dorsal view C posterior part of body, dorsal view D posterior part of body, ventral view. Scale bars: 1 mm.
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Yunnan and Sichuan karst regions of southwestern China. Given this, it is not sur-
prising to discover the genus Pacidesmus in northwestern Vietnam. The distance
between Pacidesmus tuachua sp. nov. and Pacidesmus trilobatus Liu & Golovatch,
2020 from Guangxi Province, China, is about 150 km northeast–southwest (Fig. 1).
Remarks. While there remains a noticeable geographical gap between the
mountainous northern Thailand species and the troglobionts of southern
Figure 13. Pacidesmus tuachua sp. nov., holotype ♂ (IEBR-Myr 951) right gonopod, under UV light A mesal view B ventral
view C lateral view D dorsal view. Scale bars: 1 mm.
A B
C D
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gap. Like the more eastern species, the new species is also troglobiotic, and
restricted Pacidesmus (including only the type species) needs exploration.
Pacidesmus in Vietnam.
Identication key to species of the genus Pacidesmus Golovatch, 1991
1 Sternal cones between ♂ legs 6 and 7 for accommodation of distal parts
of gonopods present. Epigean and high-montane from northern Thailand
......................................................................................................... P. shelleyi
northern Vietnam...........................................................................................2
2 Gonopod exomere absent ............................................................................3
– Gonopod exomere present ...........................................................................4
endomere rather stout, not carrying any processes. Guangxi.............P. bidus
slender, carrying two additional processes, a shorter, larger and broader
triangular one at midlength, and a longer, acuter triangular one at ¾ length.
Northern Vietnam ............................................................. P. tuachua sp. nov.
Figure 14. Pacidesmus tuachua sp. nov., holotype ♂ (IEBR-Myr 951) right gonopod A mesal view B lateral view. Abbrevia-
tions: co = coxite; pref = prefemorite; ca = cannula; en = endomere; pu = puvillus; p1p2 = second process.
Scale bars: 0.5 mm.
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
...............5
............................................................................................................... 6
5 Gonopod endomere less stout. Body length 23–24 mm, width of midbody
pro- and metazona 1.5–1.6 and 2.5–2.7 mm, respectively. Paraterga up-
turned above dorsum only on rings 1–5. Guangxi ...................... P. bedosae
– Gonopod endomere stouter. Body length 28–30 mm, width of midbody
pro- and metazona 1.4–1.7 and 2.8–3.2 mm, respectively. Paraterga up-
turned above dorsum until ring 17 (♂) or 14 (♀). Guizhou .... P. superdraco
..... 7
...............................8
7 Only paraterga 1–4 evidently upturned above dorsum. Gonopod endomere
slender, subfalcate, carrying a small tooth distally on median surface; tip
......................................................................................................P. tiani
– Paraterga upturned above dorsum starting with paraterga 2. Gonopod en-
...... P. tridus
8 Endomere with two teeth ..............................................................................9
– Endomere with either one tooth or three teeth ..........................................10
9 Caudolateral corners of paraterga strongly triangular. Gonopod exomere
.......................................................................P. whitteni
– Caudolateral corners of paraterga narrowly rounded to pointed. Gonopod
exomere smaller and unciform .................................................... P. armatus
10 Endomere with one tooth ............................................................................11
– Endomere with three teeth ..........................................................................12
11 All paraterga clearly upturned above dorsum. Endomere rather long and
strongly twisted .............................................................................. P. uncatus
– Only anterior paraterga upturned above dorsum. Endomere much shorter
and subfalcate ............................................................................... P. sinensis
base. Exomere with a large membranous process at base....... P. martensi
– Endomere long and slender, carrying three lobes. Exomere with a short
spiniform process at base .......................................................... P. trilobatus
Discussion
Beyond southern China and Southeast Asia, there are eight macropolydesmid
genera occurring in Asia, mainly in Central Asia. The main differences between
those genera are presented in Table 2 below.
Pacidesmus tuachua sp. nov. is distinguished from members of Nipponesmus
by having a gonopod endomere; the gonopod telopodite has neither a comb of
setae nor slender teeth (vs without endomere, with a conspicuous comb of setae
or slender teeth) (Golovatch et al. 2011). The new species differs from members
of Gleninea in the absence of a gonopod exomere, the gonopod endomere is with
only two additional processes, the absence of an accessory seminal chamber
(vs the presence of a gonopod exomere, the distal part of the endomere carrying
numerous or several spine-like hairs or strong, sometime curved spines, and the
presence of an accessory seminal chamber) (Golovatch and Geoffroy 2014).
The new species is also distinguished from Schizoturanius and Unira-
midesmus species by its larger size (16.3 mm vs less than 10.0 mm). The new
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
species could possibly be assigned to the genus Uniramidesmus based on
the simple, slender, falcate gonopod; however, Uniramidesmus species are all
much smaller (<10.0 mm in length), the gonopods are strongly falcate to coiled
caudally and cross each other when in situ; the opening of the seminal groove
is subterminal to terminal on a bare to more or less pubescent pulvillus (Mikhal-
jova 2004). On the contrary, the new species is far larger in size (ca 16.3 mm
in length), the gonopods in situ are well separated from each other, and the
opening of the seminal groove is on a typical hairy pulvillus.
Compared to Schizoturanius (Mikhaljova 2004), Pacidesmus tuachua sp. nov.
differs well-developed paraterga (vs a strongylosomoid, moniliform body with
narrow to almost missing paraterga, which are mostly smooth and only seldom
laterally incised); also, the gonopod is without an accessory seminal chamber (vs
with an accessory seminal chamber) and there is no gonopod femoral process
(vs with a characteristic femoral process). Finally, the new species can hardly
be assigned to the genus Jaxartes
Table 2. Morphological diagnoses and distribution of all eight macropolydesmid genera in Asia.
No. Genus Diagnosis Distribution
1Schizoturanius Body small, strongylosomoid (= without prominent paraterga), moniliform; paraterga narrow,
chamber present; gonopod femorite carrying a characteristic process (Mikhaljova 2004)
Ten species in Central Asia and
Ukraine, Asian part of Russia,
Kazakhstan, northwestern China
(Mikhaljova 2004; Nefediev 2023)
2Uniramidesmus Body small (usually ca 10 mm long); head covered with dense minute hairs; antennomeres
medium-sized and set below dorsum, with marginal incisions; metatergal polygonal sculpture
ranging from well-developed to poorly-developed; metatergal setae pointed; sphaerotrichs
present or absent; gonopods slender, strongly falcate to coiled caudally, relatively simple, in
situ crossing each other; seminal groove with a loop parabasally; accessory seminal chamber
absent; opening of seminal groove subterminally to terminally on a bare to more or less
pubescent pulvillus (Mikhaljova 2004, 2017)
Ten species in central Asia and Asian
part of Russia (Mikhaljova 2004, 2017)
3Jaxartes Body small (usually ca 10 mm long); metaterga with bosses/tubercles with bacilliform
or trichoid setae; paraterga clearly incised laterally; four distal ♂ podomeres with ventral
sphaerotrichomes; gonopod coxite without outgrowths other than cannula; the gonotelopodite
particularly slender, suberect, with the endomere being considerably longer than a ventrally
fringed process (if present at all), also bearing a parabasal tooth and a subtruncate apex, basally
et al. 2019)
4Epanerchodus Gonopod endomere mostly absent, rarely present as only a more or less rudimentary structure,
while the seminal groove after the recurvature point still makes a long way basad to debauch
into a prominent, simple-haired, accessory seminal chamber placed at the bottom of a profound
parabasal cavity in the telopodite (Liu and Golovatch 2018; Golovatch 2021).
distribution, mainly from Japan (East
Asia) to the western part of China, from
Mongolia (Central Asia) in the north to
southern China and the Himalaya of
Nepal in the south (Liu and Golovatch
2018; Golovatch 2021)
5Pacidesmus Body large, up to 30 mm long. Paraterga broad, slightly incised laterally. Metaterga with bosses
and rather simple; exomere absent or supplied with an outgrowth (Golovatch et al. 2010; Liu
and Golovatch 2020)
Twelve species in southern China
and northern part of Southeast
Asia (Golovatch et al. 2010; Liu and
Golovatch 2020)
6Nipponesmus Body size large (up to 20 mm in length). Paraterga broad. Gonopod endomere with conspicuous
comb of setae or slender teeth. The seminal groove running mostly mesally to recurve neatly
between exomere and endomere, then to debauch somewhat basally into a prominent hairy
pulvillus which also beset with the same peculiar trichome, and is devoid of an accessory
seminal chamber (Golovatch et al. 2011)
Three species in Japan and Taiwan.
(Golovatch et al. 2011)
7Gleninea Body small to large size (up to 16 mm in length). The third pair of ♂ legs only slightly thickened.
Antenomere 5-6, each with a s mall, compact, distodorsal group of bacilliform sensilla.
Lateral side of paraterga strongly serrated, 5-6 small, sharp teeth. Gonopod exomere simple,
subfalcate; endomere with distal part carrying numerous or several spine-like hairs or strong,
sometime curved spines. Accessory seminal chamber present (Golovatch and Geoffroy 2014)
Seven species in the Himalaya of India,
Nepal, Bhutan, and China (Golovatch
and Geoffroy 2014)
8Polydesmus Body size medium to large. Paraterga usually wide. Gonopod solenomere absent to rather well
developed, sometimes conspicuously shaped. Exomere from short and slightly curved to very
long and strongly falcate, mostly uniramous, directed caudally, starting laterally or apically and
basally of recurvature point of seminal groove. Seminal groove, largely mesal; terminal laterad
loop relatively short and turning around a distofemoral process (Djursvoll et al. 2001)
About 200 species distributed mainly
in the Mediterranean; a few species
in East Asia and northern Vietnam
(Djursvoll et al. 2001)
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
larger body size (16 mm long), the slender, strongly falcate gonopods without an
accessory seminal chamber. On the contrary, the genus Jaxartes is diagnosed by
its small body (usually ca 1 cm long); the metaterga show bosses or tubercles
with bacilliform or trichoid setae, the paraterga are clearly laterally incised, there
are four distal male podomeres with ventral sphaerotrichomes, the gonopod cox-
ite is without outgrowths apart from the typical cannula, and the gonotelopodite
is particularly slender, suberect, and with the endomere being considerably lon-
ger than a ventrally fringed process (if present at all); also, the endomere bears
a parabasal tooth and has a subtruncate apex, basally with a very evident hairy
Pacidesmus tuachua sp. nov. can hardly be placed in Epanerchodus or
Polydesmus because its paraterga are relatively narrow, the seminal groove
starts mesally, as usual, then is recurved laterad at the base of a particular-
ly prominent endomere branch to enter an accessory seminal chamber that
opens on a setose pulvillus, and the endomere bears additional processes.
The strongly sigmoid gonopodal telopodite in P. tuachua sp. nov is somewhat
unusual in comparison to that in other Pacidesmus species. This difference may
suggest a new genus; however, it currently seems best assigned to Pacidesmus
based on the above discussion. It is noteworthy that most Schizoturanius or
Polydesmus spp. likewise show only slightly curved gonopod telopodites, but
relatively few species in these genera are so strongly sigmoid.
To support further study of Polydesmidae in Vietnam and Southeast Asia,
An identication key to macropolydesmid genera in Asia
Based on Turk (1945), Golovatch (1991), Djursvoll et al. (2001), Mikhaljova (2004),
1 Gonopods without an accessory seminal chamber (Uniramidesmus, Nip-
ponesmus, Jaxartes) ..................................................................................... 2
– Gonopods with an accessory seminal chamber (Schizoturanius, Pac-
idesmus, Gleninea, Polydesmus, Epanerchodus) .........................................4
2 Body moniliform, paraterga narrower. Seminal groove running mostly
mesally to recurve neatly between exomere and endomere. Gonopod en-
.............Nipponesmus
– Body not moniliform, paraterga broader. Seminal groove running entirely me-
... 3
3 Seminal groove opening subterminally to terminally on a bare to more or
less pubescent pulvillus ........................................................Uniramidesmus
– Seminal groove opening on a distinct, ventral, hairy pulvillus ....... Jaxartes
4 Body moniliform, paraterga narrow. Loop of seminal groove distal ............
................................................................................................. Schizoturanius
– Body not moniliform, paraterga broad. Loop of seminal groove not distal .... 5
5 Paraterga strongly serrated or incised laterally. Gonopod endomere in dis-
tal part carrying numerous or several spine-like hairs or strong, sometime
curved spines.................................................................................... Gleninea
– Paraterga not strongly serrated/incised laterally. Gonopod endomere without
numerous, mostly strong, sometimes curved spines or bacilli or setae......... 6
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
6 Paraterga narrower. Seminal groove starting mesally, then recurving lat-
erad at the base of a particularly prominent endomere branch to enter an
accessory seminal chamber that opens on a setose pulvillus; endomere
carrying additional processes ................................................... Pacidesmus
– Paraterga wider. Seminal groove largely running mesally. Gonopod en-
domere absent ............................................................................................... 7
7 An endomere absent. Exomere from short and slightly curved to very long
and strongly falcate, mostly uniramous, directed caudally, starting laterally or
apically and basally of recurvature point of seminal groove ......... Polydesmus
– An endomere mostly absent, but rather rarely present as only a more
or less rudimentary structure, while the seminal groove after the recur-
vature point still makes a long way basad to debauch into a prominent,
simple-haired, accessory seminal chamber placed at the bottom of a pro-
found parabasal cavity in the telopodite ................................Epanerchodus
Conclusion
Two polydesmid genera and species are presently known to occur in Vietnam:
Polydesmus vietnamicus Nguyen, 2009 and Pacidesmus tuachua sp. nov.
Polydesmus vietnamicus is an epigean, forest-dwelling species, while Pacidesmus
tuachua is troglobiotic. The diversity of polydesmids in Vietnam is potentially
greater given the number of species in other southern Asian regions. The
paucity of species is either due to some yet-unknown historical, evolutionary
surveys are needed to more fully clarify the diversity and biogeography of
polydesmids in Southeast Asia.
Acknowledgements
We thank Dang Van Dong and Nguyen Duc Hiep, from IEBR, for their help in
USA), Dr Sergei I. Golovatch (Russian Academy of Sciences, Moscow, Russia)
and Dr Natdanai Likhitrakarn (Faculty of Agricultural Production, Maejo Univer-
sity, Chiang Mai, Thailand) are acknowledged for kindly checking, revising, and
correcting the submission. The work is supported by the Vietnam Academy of
Science and Technology under the project NCXS01.04/23-25.
Additional information
Conict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
The work is funded by the Vietnam Academy of Science and Technology under the proj-
-
versity and application potential of hymenopterans, myriapods and soil nematodes in
278
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Anh D. Nguyen et al.: Polydesmid millipedes from Vietnam
the limestone mountains of northeastern Vietnam”. This study has also been supported
by the following funds: the Fund for the Promotion of Joint International Research
(Fostering Joint International Research (B)) (JSPS KAKENHI, no. 22KK0087, Leader:
23K05299, Leader: E Oguri, FY2023-2026), the Tokyo Metropolitan University Fund for
TMU Strategic Research (leader: Prof. Noriaki Murakami, FY2020–FY2022), and the
Asahi Glass Foundation (leader: K. Eguchi, FY2017–FY2022).
Author contributions
Conceptualization: ADN. Data curation: ADN, KE. Formal analysis: ADN. Funding acqui-
sition: KE, TTTV. Resources: TTTV. Visualization: TTTV. Writing - original draft: ADDN.
Writing - review and editing: KE, ADN.
Author ORCIDs
Anh D. Nguyen https://orcid.org/0000-0001-9273-0040
Tam T. T. Vu https://orcid.org/0000-0003-1145-975X
Katsuyuki Eguchi https://orcid.org/0000-0002-1054-1295
Data availability
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