Content uploaded by Ambalaparambil V. Sudhikumar
Author content
All content in this area was uploaded by Ambalaparambil V. Sudhikumar on Jun 09, 2021
Content may be subject to copyright.
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by T. Wesener: 28 Apr. 2021; published: 1 Jun. 2021 373
Zootaxa 4980 (2): 373–382
https://www.mapress.com/j/zt/
Copyright © 2021 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4980.2.8
http://zoobank.org/urn:lsid:zoobank.org:pub:C55A445F-76EC-460A-B259-98AF744ECFFF
The millipede genus Klimakodesmus Carl, 1932, with the description of a new species
from Kerala state, southern India (Diplopoda, Polydesmida, Pyrgodesmidae)
MATHILAKATH DASAN ASWATHY1,3, SERGEI I. GOLOVATCH 2* &
AMBALAPARAMBIL VASU SUDHIKUMAR1,4
1Centre for Animal Taxonomy and Ecology (CATE), Christ College, Irinjalakuda, Kerala, India
2Institute for Problems of Ecology and Evolution, Russian Academy of Sciences,
Leninsky pr. 33, Moscow 119071, Russia
3
�
aswathym.das94@gmail.com; https://orcid.org/0000-0002-3904-8066
4
�
spidersudhi@gmail.com; https://orcid.org/0000-0002-4479-4995
*Corresponding author.
�
sgolovatch@yandex.ru; https://orcid.org/0000-0001-7159-5484
Abstract
Klimakodesmus Carl, 1932 is briefly redescribed, rediagnosed, and shown to be an oligotypic genus endemic to southern
India and distinct from the particularly similar genus Pyrgodesmus Pocock, 1892, monobasic and endemic to Sri
Lanka, by several important features of peripheral and, especially, gonopodal structure. A new species, Klimakodesmus
bilobocaudatus sp. nov., is described from Kerala state, India, differing from the sole accepted, and type species K.
gravelyi Carl, 1932, from Tamil Nadu state, primarily by the laterally trilobate paraterga, the caudally more deeply
bilobate mid-dorsal keel on ring 19, and certain minor details of the gonopodal structure.
Key words: Klimakodesmus, Pyrgodesmus, taxonomy, new species, India, Sri Lanka
Introduction
At present, the millipede fauna of India, however insufficiently studied, comprises 270+ nominate species or sub-
species in at least 90 genera, 25 families and 11 orders (Golovatch & Wesener 2016). Of them, the large and mostly
tropical family Pyrgodesmidae contains some 14 species in seven genera, including Klimakodesmus Carl, 1932.
The taxonomy of Pyrgodesmidae is long known to be particularly badly confused and chaotic globally, with
more than 170 genera, largely monotypic, and nearly 400 species all over the tropics, marginally in Southern Eu-
rope, North Africa, southern U.S.A., Japan, Taiwan and central China (Enghoff et al. 2015). Progress is strongly
hampered by the small size of the animals (typically 3–16 mm long), as well as the outstandingly diverse and often
complex structures, both peripheral and gonopodal, which are often difficult to see, let alone describe and depict.
Numerous pyrgodesmid genera remain monotypic, often based only on female or even juvenile material, and delim-
ited using solely such somatic features as the number of body segments/rings, the shape, distribution and location
of ozopores and tergal ornamentations, the structure of the paraterga etc.
Klimakodesmus was originally described based on a single, and type species, K. gravelyi Carl, 1932, originating
from near Coonoor and Mudumalai, both in Tamil Nadu state, Western Ghats, southern India (Carl 1932). Verhoeff
(1939) promoted Klimakodesmus to the rank of a monobasic subfamily, Klimakodesminae Verhoeff, 1939, but
presently it is considered as a strict junior subjective synonym of Pyrgodesmidae (Hoffman 1980; https://www.
fieldmuseum.org/sites/default/files/taxon_table_5.pdf).
Carl (1932), when erecting his Klimakodesmus, compared it to Pyrgodesmus Pocock, 1892, the type genus of
Pyrgodesmidae and also monobasic (Pocock 1892), because both genera shared numerous somatic features. He paid
special attention to the identity of Pyrgodesmus obscurus Pocock, 1892, the type species of Pyrgodesmus, described
from two male syntypes from Pundaluoya, Sri Lanka. Carl (1932) compared the original, rather superficial descrip-
tion and few schematic drawings of P. obscurus as given by Pocock (1892) to the excellent and richly illustrated
ASWATHY ET AL.
374 · Zootaxa 4980 (2) © 2021 Magnolia Press
redescription of that species as presented by Silvestri (1920), based on a male syntype from Pundaluoya and two
additional males from Paradenyia, Sri Lanka. Carl noted differences between both descriptions in tergal ornamenta-
tion, antennae, leg thickness, and gonopodal conformation. In particular, the gonopods of P. obscurus were verbally
described as showing unusually hypertrophied coxites and blade-like telopodites (Pocock 1892), vs. moderately
enlarged and squarish coxites with strongly curved and ribbon-shaped telopodites as depicted by Silvestri (1920).
As a result, Carl (1932) considered Silvestri’s (1920) record and most detailed redescription of P. obscurus as a
misidentification, suggesting to give it a new name different from P. obscurus, and emphasizing that neither spe-
cies belonged to Klimakodesmus. He also expressed the need for a new revision of the P. obscurus syntypes, both
presumably kept in the London Museum (Pocock 1892).
On the other hand, according to Silvestri (1920), the tegument of at least some of the Pyrgodesmus obscurus
samples he studied was so heavily coated with earth that only after it had been thoroughly cleansed did it show the
necessary details of the underlying, real tergal structure and sculpture. Moreover, both bodies, one coated with earth
and the other clean, were very beautifully and artistically illustrated, albeit neither was properly labeled as to its
provenance! So we can only adhere to Carl’s (1932) appeal to revise at least one of the male syntypes of P. obscurus,
since the task is to simply check the gonopodal structure and to compare it to Silvestri’s (1920) account and illustra-
tions. Only this could finally reveal the species’ identity.
It was Attems (1940) who took action following Carl’s (1932) investigation and suggestions. He proposed Kli-
makodesmus permutatus Attems, 1940 as a replacement name for Pyrgodesmus obscurus in the sense of Silvestri
(1920), but, contrary to Carl’s opinion, he assigned it to Klimakodesmus. It is since then that formally the genus
Klimakodesmus has hitherto contained two species: K. gravelyi Carl, 1932, from southern India (Golovatch & We-
sener 2016), and K. permutatus Attems, 1940, from Sri Lanka (De Zoysa et al. 2016). Moreover, strangely enough,
Attems (1940: 271) retypified Klimakodesmus through selecting K. permutatus Attems, 1940 as the type species, a
completely invalid action (Jeekel 1971).
The present paper puts on record a second, new species of Klimakodesmus, also clarifying the similarities to
and differences from Pyrgodesmus. We follow Carl (1932) in treating both these genera as distinct and, pending a
revision of P. obscurus, we remove K. permutatus from the scope of Klimakodesmus (see the reasons in Carl (1932)
and below).
Materials and methods
The material underlying the present contribution was hand-collected in a sacred grove of Kannur district, Kerala, In-
dia. The local name of the grove is Thekkumbadu Koolom—Thazhekkavu, located on an inland island bordered by
thick mangroves. The samples were collected in January 2021 (winter season) and preserved in absolute ethanol.
The holotype and most of the paratypes are deposited in the collection of the Centre for Animal Taxonomy and
Ecology, Christ College, Irinjalakuda, Kerala, India (CATE), with the holotype ID–CATE-5302, and paratypes
ID–CATE-5302B–CATE-5302I. A few paratypes have been lent for further studies to the Zoological Museum, State
University of Moscow (ZMUM), Russia, as indicated below.
Photographs in the field were taken with a Canon EOS 5D Mark-III camera using a Canon EF 100 mm f/2.8
Macro USM Lens and a Canon MT-24EX Macro Twin Lite Flash. Multifocal photographs of specimens were taken
in the lab with a Leica DMC4500 digital camera mounted on a Leica M205 C stereo microscope. Photographs were
stacked, and the measurements (in mm) taken, using Leica Application Suite (LAS) version 4.3.0 software. The
final images were processed with Adobe Photoshop CC software. The map was executed using both Google Earth
and Simple Mappr software.
In the following morphological accounts, we use Hoffman’s (1976) formula for describing the most typical
ornamentation pattern of the metaterga in Pyrgodesmidae, in which:
PM stands for paramedian,
DL for dorsolateral tubercles/lobes,
i for intercalary series of grains,
Am for anteromarginal,
Cm for caudomarginal lobulations.
THE MILLIPEDE GENUS KLIMAKODESMUS Zootaxa 4980 (2) © 2021 Magnolia Press · 375
Taxonomic part
Order Polydesmida Latreille, 1802/03
Family Pyrgodesmidae Silvestri, 1896
Genus Klimakodesmus Carl, 1932
Type-species: Klimakodesmus gravelyi Carl, 1932, by monotypy.
Other species included: Klimakodesmus bilobocaudatus sp. nov.
Brief description. This genus shows a body shape and an ornamentation pattern typical of Pyrgodesmidae, charac-
terized by 20 segments/rings (19+T) both in the male and female; a flabellate collum is strongly domed and tubercu-
late, completely covering the head from above, with 5+5 distinct, equal, clearly incised and rounded lobulations at a
subhorizontal anterior margin; the antennae are short, two basal antennomeres of each antenna being sunken inside a
distinct transverse groove, and antennomere 5 being much longer and thicker than the 6th; a cerotegument crust with
microvilli covers most of the dorsal surface of the collum and following metaterga, the limbus being microlobulate
and microspiculate caudally; body rings are strongly arched dorsally, each with 2+2 longitudinal rows of basically
2+2 or 3+3 larger and fused tubercles/lobes (paramedian, PM, and dorsolateral, DL), both rows largely representing
higher bilobed keels mostly slightly inclined either forward or caudad, but PM always being higher than DL. PM of
only ring 19 is abruptly and unusually strongly elongate and more or less clearly bilobate caudally, conspicuously
overhanging and concealing a short epiproct from above (Figs 2–4, 8–10). Neither intercalary (i) series of grains nor
antero- (Am), nor truly caudomarginal lobulations (Cm) are distinguished. The pore formula is normal: ozopores 5,
7, 9, 10, 12, 13, 15 and 16 are borne on distinct porosteles, vs. 17, 18 and 19 which open flush on the surface. The
paraterga are set low (at about half the height of midbody metazonae), slightly declivous to subhorizontal, leaving
the dorsum very convex, moderately bi- or trilobate laterally regardless of a cylindrical porostele or ozopore located
just before or upon the ultimate lobulation, respectively. The legs are short and stout, rather sparsely setose, the
prefemur bearing a particularly strong seta distomesally, the tibia a distodorsal one.
The gonopods are fairly simple, in situ either slightly diverging or distally crossing each other; each coxite is
moderately enlarged, hemispherical or squarish, microgranulate and microsetose laterally, and concave medially,
the gonocoel thus being moderately deep; the cannula is as usual, simple, moderately long and strongly unciform.
The telopodite is simple, slender, unipartite, represented by a very strong, moderately long, but mostly exposed,
gradually attenuating and acuminate solenomere.
Diagnosis. Klimakodesmus seems to be particularly similar to Pyrgodesmus in most somatic characters, as
correctly noted by Carl (1932), but both genera are distinct primarily in gonopodal structure: clearly hypertrophied
gonocoxites and “blade-like” telopodites (Pocock 1892) vs. moderately incrassate, apparently smaller, subspherical
or squarish gonocoxites, each of which supports a strong and unipartite telopodite represented solely by a slender,
strongly exposed, increasingly attenuating and acuminate solenomere. In addition, PM crests, including the one
on the penultimate ring that overhangs and fully conceals the epiproct from above, are much higher and stronger,
whereas DL reduced, in Pyrgodesmus compared to Klimakodesmus species.
A similar, remarkably strong, massive and largely exposed solenomere taking up most of the gonopod telopodite
is also observed in the genus Pseudocatapyrgodesmus Miyosi, 1957, with P. glaucus Miyosi, 1957, the type species
from Honshu, Japan (Miyosi 1957), and P. pulcher Golovatch, Semenyuk, VandenSpiegel & Anichkin. 2011, from
southern Vietnam (Golovatch et al. 2011). Yet their gonopods are considerably more elaborate and clearly branched,
vs. relatively very simple and unipartite in Klimakodesmus. They are the gonopods of P. pulcher that seem to be
especially similar to those of Klimakodesmus (Golovatch et al. 2011).
The strong, flattened, dorsal crest PM on the penultimate ring that overhangs and conceals the epiproct from
above, however conspicuous, is unique to neither Klimakodesmus nor Pyrgodesmus species. For instance, several,
but not all of the Afrotropical species of the genera Monachodesmus Silvestri, 1927 or Udodesmus Cook, 1896 show
the same or very similar conditions (Golovatch et al. 2015, 2017).
ASWATHY ET AL.
376 · Zootaxa 4980 (2) © 2021 Magnolia Press
Species description
Klimakodesmus bilobocaudatus sp. nov.
Figs 1–15
Material examined. Adult male holotype (CATE-5302), 2 adult male and 6 subadult female paratypes (CATE-
5302B–CATE-5302I), 1 adult male and 2 subadult female paratypes (ZMUM), India, Kerala state, Kannur district,
N11°58’2.5”, E75°17’46.6”, 5 m a.s.l., 27/01/2021, M.D. Aswathy leg.
Name. To emphasize a caudally relatively deeply bilobed dorsal keel (PM) on ring 19 in the new species; adjec-
tive.
Diagnosis. The new species differs clearly from K. gravelyi, the only other congener to be accepted (Carl 1932),
by the more regularly tuberculate collum (cf. Figs 5 and 16), the lower PM and DL which are mostly inclined caudad
(vs. inclined forward until ring 16, cf. Figs 7, 8 and 17), the caudally more deeply bilobed dorsal keel (PM) on ring
19 (vs. indistinctly bilobulate, cf. Figs 9, 10 and 18), the laterally bi- (rings 3 and 4) or trilobate (remaining postcol-
lum rings) paraterga (vs. mostly bilobate, trilobate only on ring 2, cf. Figs 5, 8, 9 and 17–19), and the gonopods
showing subspherical coxites (vs. squarish) and subcontiguous, apically subunciform and crossing each other (vs.
divergent and suberect) telopodites, each of the latter additionally reinforced with a mesobasally thickened wall (vs.
a laterobasal lobe, cf. Figs 12–15 and 20 & 21).
Description. Length ca. 3.68 mm, width of midbody segments 0.10 and 0.14 mm on pro- and metazonae, re-
spectively (holotype). Length of adult male paratypes ca. 3.0–3.8 mm, that of subadult female paratypes 3.2–3.5
mm. Width of adult male paratypes ca. 0.40–0.45 mm, that of subadult female paratypes 0.44–0.48 mm. Live co-
louration generally pink; antennae, legs and venter translucent, lighter than dorsum (Figs 1, 7 & 9). Colouration in
alcohol after 2 months of preservation pale brown to pinkish brown, legs and antennae faded to translucent (Figs
2–6, 8 & 10). No earth crust visible on body.
Most characters as in the generic description above, except as follows.
Body with 20 segments (likely in both sexes, see below) (Figs 2 & 4). In width, head << collum < segment
2=3 < 4–6 < 7–9 < 10–15, body gradually tapering towards telson thereafter. Head circular, labral region moder-
ately setose, epicranial suture rather deep, vertex microgranulate (Fig. 6). Interantennal isthmus nearly as wide as
diameter of antennal socket. Antennae short, weakly clavate, in situ reaching body segment/ring 3 when stretched
ventrolaterally. In length, antennomere 7<3<1<6<4<2<<5; antennomeres 5–7 each with a more or less compact
apicodorsal group of bacilliform sensilla (Figs 6 & 7). Collum flabellate, completely covering the head from above;
anterior margin with 5+5 distinct, equal, moderately incised and rounded lobulations; central part domed, with 2+2
and 3+3 prominent and rounded tubercles arranged in two transverse rows (Figs 2–7). Paraterga set low (at about
half of midbody height), subhorizontal to faintly declivous. Dorsum strongly arched, its outline smoothly extend-
ing onto paraterga (Figs 3 & 7). Prozonae, as well as deep and thin strictures between pro- and metazonae finely
alveolate. Postcollum metaterga with strongly differentiated dorsal tuberculations: 2+2 longitudinal crests/rows of
mostly 2+2 fused tubercles, those before ring 19 slightly inclined caudad, but PM on ring 19 abruptly modified
into a mid-dorsal, strongly elongate, conspicuous and caudally clearly bilobate ridge overhanging and concealing a
short epiproct (Figs 2–4, 8–10). Paraterga broad, thin, rather irregularly tuberculate near bases and clearly lobulated
laterally, with three rounded and moderately incised lobulations on rings 2–19, regardless of cylindrical porosteles
located between the 2nd and 3rd lobulations on rings 5, 7, 9, 10, 12, 13, 15 and 16 (Figs 2–10); only paraterga 3 and
4 with last lobulation in a clearly caudolateral position (Figs 5–7). Epiproct (Fig. 10) finger-shaped, tip bent down
and divided into two round bulbs. Hypoproct (Fig. 10) semi-circular, with 1+1 strong setae borne on minute, caudal,
paramedian knobs.
Legs with neither adenostyles nor tarsal brushes (Fig. 11). In length, tarsi > femora > prefemora > tibiae > coxae
> postfemora; claws simple, slightly curved ventrally.
Gonopods (Figs 12–15) relatively simple, placed inside a transversely oblong-oval gonopodal aperture, the lat-
ter with thin and slightly elevated caudal and lateral margins; in situ, both gonopods strongly exposed, held mostly
parallel to one another, with only tips crossing each other. Coxites (cx) voluminous, but moderately enlarged, gono-
coel thus being relatively shallow; cannulae simple. Telopodites directed caudally, each unipartite and subtriangular,
long and slender, gradually attenuating distad and acuminate apically, each represented solely by a solenomere (sl)
and supporting a seminal groove running along lateral margin to the end of a laterally directed and unciform apical
THE MILLIPEDE GENUS KLIMAKODESMUS Zootaxa 4980 (2) © 2021 Magnolia Press · 377
part, and additionally reinforced basally through a distinctly thickened caudomesal wall of a prominent, membra-
nous, mesal lobe (lo); prefemorite as usual, relatively short, densely and strongly setose.
FIGURES 1–4. Klimakodesmus bilobocaudatus sp. nov. 1. Natural microhabitat (rotting wood with a colony on the surface,
photographed not to scale). 2. Adult male paratype, dorsal view. 3. Subadult female paratype, lateral view. 4. Adult male para-
type, ventral view.
ASWATHY ET AL.
378 · Zootaxa 4980 (2) © 2021 Magnolia Press
FIGURES 5–7. Klimakodesmus bilobocaudatus sp. nov. Anterior parts of body, dorsal, ventral and lateral views, respec-
tively.
Ecological notes. This new species was found under rotting wood near a lowland swamp overgrown with My-
ristica forest in the Kannur district, northern Kerala, India. The habitat area of approximately 20 ha is a sacred grove
protected by local people. In contrast, K. gravelyi came from high-altitude areas (1000–1600 m a.s.l.) in the state of
Tamil Nadu (Carl 1932), Western Ghats, southern India (Fig. 22).
It seems noteworthy that all females collected at the type locality in January appear to be subadults, each with
19 body segments. This has been proven by dissecting the vulvae which have always been found underdeveloped
(amorphous ovoid clods devoid of setae), and an anterior, ventral, transverse, marginal ridge on ring 3 missing.
This agrees well with the common wisdom that only adult males in Polydesmida, but never adult females, can be
composed of a lesser number of body rings. Usually, in the course of teloanamorphosis characteristic of the order
Polydesmida, both the males and females of Polydesmida show the same number of body rings, either 20 or 19,
THE MILLIPEDE GENUS KLIMAKODESMUS Zootaxa 4980 (2) © 2021 Magnolia Press · 379
more rarely 18, exceptionally up to 40 (Shear et al. 2016). When there is a difference in polydesmidan body seg-
ment counts, the adult males typically have one segment less and nearly always remain stable per sex (Enghoff et al.
2015). Only exceptionally, as is the case of Ammodesmus granum Cook, 1896 and A. congoensis VandenSpiegel &
Golovatch, 2015 (Ammodesmidae), from western or central Africa, respectively, can both sexes vary in the number
(18 or 19) of body rings (VandenSpiegel & Golovatch 2012, 2015).
FIGURES 8–15. Klimakodesmus bilobocaudatus sp. nov. 8. Posterior half of body (subadult female paratype), lateral view.
9 & 10. Posterior part of body (adult male paratype), dorsal and ventral views, respectively. 11. Midbody leg (adult male para-
type), lateral view. 12–14. Both gonopods (adult male paratype), caudal, oral and caudal views, respectively. 15. Right gonopod,
caudal view. Abbreviations: cx, coxite; lo, mesal lobe; sl, solenomere. NB: Figures 14 & 15 drawn not to scale.
ASWATHY ET AL.
380 · Zootaxa 4980 (2) © 2021 Magnolia Press
FIGURES 16–21. Klimakodesmus gravelyi Carl, 1932, male holotype, after Carl (1932). 16. Collum, dorsal view. 17. Anterior
part of body, lateral view. 18 & 19. Posterior part of body, dorsal and lateral views, respectively. 20 & 21. Left and right gonop-
ods, caudal and oral views, respectively. Abbreviations: cx, coxite; lo, lateral lobe; sl, solenomere. Drawn not to scale.
In Klimakodesmus species, however, the situation is ordinary, as both sexes (must) equally have 20 body seg-
ments. The absence of adult females in the type series seems best to be accounted for by seasonal factors alone.
THE MILLIPEDE GENUS KLIMAKODESMUS Zootaxa 4980 (2) © 2021 Magnolia Press · 381
FIGURE 22. Distribution of the genus Klimakodesmus: K. gravelyi in Tamil Nadu state (orange dots), and K. bilobocaudatus
sp. nov. in Kerala state (red dot).
Acknowledgements
The authors are grateful to the Principal, Christ College (Autonomous), Irinjalakuda, Kerala, for providing us with
the facilities necessary for undertaking this study. The first author is thankful to UGC-JRF for the financial sup-
port to conduct this study. The authors also acknowledge the funding rendered by Science & Engineering Research
Board (SERB), DST for the facilities used in this study (Major Research Project EMR/2016/006401). The second
author was partly supported by the Presidium of the Russian Academy of Sciences, Program No. 41 “Biodiversity
of Natural Systems and Biological Resources of Russia”. We are also grateful to Mr. Santhosh, Devaswom staff, and
Mr. Chandran, care taker of the sacred grove, for their hospitality and support in the field. We are greatly obliged
to Nesrine Akkari (Vienna, Austria), Didier VandenSpiegel (Tervuren, Belgium) and Pooja A. Anilkumar (Bonn,
Germany), the three reviewers whose critiques and valuable suggestions have allowed us to considerably improve
the paper.
References
Attems, C. (1940) Myriopoda 3. Polydesmoidea III. Fam. Polydesmidae, Vanhoeffeniidae, Cryptodesmidae, Oniscodesmidae, Sphaeri-
otrichopidae, Peridontodesmidae, Rhachidesmidae, Macellolophidae, Pandirodesmidae. Das Tierreich, 70, i–xxxii + 1–577.
Carl, J. (1932) Diplopoden aus Süd-Indien und Ceylon. 1. Teil. Polydesmoidea. Revue suisse de Zoologie, 39 (17), 411–529.
https://doi.org/10.5962/bhl.part.118948
De Zoysa, H.K.S., Nguyen, D.A. & Wickramasinghe, S. (2016) Annotated checklist of millipedes (Myriapoda: Diplopoda) of
Sri Lanka. Zootaxa, 4061 (5), 451–482.
https://doi.org/10.11646/zootaxa.4061.5.1
ASWATHY ET AL.
382 · Zootaxa 4980 (2) © 2021 Magnolia Press
Enghoff, H., Golovatch, S.I., Short, M., Stoev, P. & Wesener, T. (2015) Diplopoda – taxonomic overview. In: Minelli, A. (Ed.)
Treatise on Zoology—Anatomy, Taxonomy, Biology. The Myriapoda. 2 (16). Brill, Leiden and Boston, pp. 363–453.
https://doi.org/10.1163/9789004188273_017
Golovatch, S.I. & Wesener, T. (2016) A species checklist of the millipedes (Myriapoda, Diplopoda) of India. Zootaxa, 4129 (1),
1–75.
https://doi.org/10.11646/zootaxa.4129.1.1
Golovatch, S.I., Nzoko Fiemapong, A.R. & VandenSpiegel, D. (2015) Notes on Afrotropical Pyrgodesmidae, 2 (Diplopoda:
Polydesmida). Arthropoda Selecta, 24 (4), 387–400. [https://kmkjournals.com/upload/PDF/ArthropodaSelecta/24/24_4_
387_400_Golovatch_et_al_for_Inet.pdf]
https://doi.org/10.15298/arthsel.24.4.02
Golovatch, S.I., Nzoko Fiemapong, A.R. & VandenSpiegel, D. (2017) Notes on Afrotropical Pyrgodesmidae, 3 (Diplopoda:
Polydesmida). Arthropoda Selecta, 26 (3), 175–215. [https://kmkjournals.com/upload/PDF/ArthropodaSelecta/26/26_3_
175_215_Golovatch_et_al_for_Inet.pdf]
https://doi.org/10.15298/arthsel.26.3.01
Golovatch, S.I., Semenyuk, I.I., VandenSpiegel, D. & Anichkin, A.E. (2011) Three new species of the millipede family Pyrgo-
desmidae from Nam Cat Tien National Park, southern Vietnam (Diplopoda: Polydesmida). Arthropoda Selecta, 20 (1), 1–9.
[https://kmkjournals.com/upload/PDF/ArthropodaSelecta/20/20_1%20001_009%20Golovatch.pdf]
https://doi.org/10.15298/arthsel.20.1.01
Hoffman, R.L. (1976) A new lophodesmid milliped from a Guatemalan cave, with notes on related forms (Diplopoda: Pyrgo-
desmidae). Revue suisse de Zoologie, 83 (2), 307–316. [https://archive.org/details/biostor-126721]
https://doi.org/10.5962/bhl.part.91441
Hoffman, R.L. (1980) Classification of the Diplopoda. Muséum d’histoire naturelle, Genève, 237 pp.
Jeekel, C.A.W. (1971) Nomenclator generum et familiarum Diplopodorum: A list of the genus and family-group names in the
Class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Monografieёn van de Nederlandse Entomolo-
gische Veneniging, 5, i–xii + 1–412. [for 1970]
Miyosi, Y. (1957) Beiträge zur Kenntnis japanischer Myriopoden. 20. Aufsatz: Über eine neue Gattung von Diplopoda, eine
neue Art und eine neue Unterart von Chilopoda. Zoological Magazine, Tokyo, 66 (6), 264–268. [in Japanese, with English
abstract]
Pocock, R.I. (1892) Report upon two collections of Myriapoda sent from Ceylon by Mr. E. E. Green, and from various parts of
southern India by Mr. Edgar Thurston, of the Government Central Museum, Madras. Journal of the Bombay Natural His-
tory Society, 7, 131–174. [https://www.biodiversitylibrary.org/item/95456#page/7/mode/1up]
Shear, W.A., Ferreira, R.L., Iniesta, L.F.M. & Marek, P. (2016) A millipede missing link: Dobrodesmidae, a remarkable new
polydesmidan millipede family from Brazil with supernumerary rings (Diplopoda, Polydesmida), and the establishment of
a new suborder Dobrodesmidea. Zootaxa, 4178 (3), 371–390.
https://doi.org/10.11646/zootaxa.4178.3.4
Silvestri, F. (1920) Descriptions of some Oriental Diplopoda Polydesmoidea of the subfamily Pyrgodesminae. Records of the
Indian Museum, 19 (4), 117–135.
VandenSpiegel, D. & Golovatch, S.I. (2012) The millipede family Ammodesmidae (Diplopoda, Polydesmida) in western Africa.
ZooKeys, 221, 1–17.
https://doi.org/10.3897/zookeys.221.3739
VandenSpiegel, D. & Golovatch, S.I. (2015) A new millipede of the family Ammodesmidae found in central Africa (Diplopoda,
Polydesmida). ZooKeys, 483, 1–7.
https://doi.org/10.3897/zookeys.483.9150
Verhoeff, K.W. (1939) Diplopoden der Insel Mauritius und ihre zoogeographische Bedeutung. Jenaische Zeitschrift für Natur-
wissenschaft, 73, 37–96.
