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Greater Hog Badger Arctonyx collaris (Mustelidae, Carnivora) from the Pleistocene of Vietnam (Lang Trang Cave)

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  • Borissiak Paleontological Institute of the Russian Academy of Sciences

Abstract and Figures

The dental remains of a large badger collected in 2020 from the Pleistocene deposits of the Lang Trang karstic cave in northern Vietnam are described. The molars М 1 and М 1 exhibit the features that are characteristic of Arctonyx collaris Cuvier, 1825. At the same time, the length of M 1 of the Lang Trang form exceeds that of modern greater hog badgers. Based on the size and structure of the teeth, it can be assigned to the subspecies A. c. rostratus Matthew et Granger, 1923 known from the Pleistocene of China and Southeast Asia.
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ISSN 0012-4966, Doklady Biological Sciences, 2020, Vol. 495, pp. 280–282. © Pleiades Publishing, Ltd., 2020.
Russian Text © The Author(s), 2020, published in Doklady Rossiiskoi Akademii Nauk. Nauki o Zhizni, 2020, Vol. 495, pp. 577–580.
Greater Hog Badger Arctonyx collaris (Mustelidae, Carnivora)
from the Pleistocene of Vietnam (Lang Trang Cave)
Academician A. V. Lopatina,*
Received June 23, 2020; revised July 10, 2020; accepted July 10, 2020
AbstractThe dental remains of a large badger collected in 2020 from the Pleistocene deposits of the Lang
Trang karstic cave in northern Vietnam are described. The molars М1 and М1 exhibit the features that are
characteristic of Arctonyx collaris Cuvier, 1825. At the same time, the length of M1 of the Lang Trang form
exceeds that of modern greater hog badgers. Based on the size and structure of the teeth, it can be assigned to
the subspecies A. c. rostratus Matthew et Granger, 1923 known from the Pleistocene of China and Southeast
Asia.
Keywords: Arctonyx collaris, Melinae, Mustelidae, Carnivora, greater hog badger, Pleistocene, Lang Trang
cave, Vietnam
DOI: 10.1134/S0012496620060058
The greater hog badger Arctonyx collaris Cuvier,
1825 is one of the largest extant badgers (with a condy-
lobasal skull length up to 172–174 mm, body length up
to 87–88 cm, weight up to 14–21.5 kg) [1–3]. The
species currently inhabits eastern India and Indochina
reaching peninsular Myanmar and Thailand [1, 3]. In
Middle to Late Pleistocene, A. collaris was a common
member of the StegodonAiluropoda fauna of China
and Southeast Asia; with the fossil forms usually
assigned to the extinct subspecies A. collaris rostratus
Matthew et Granger, 1923 [1, 4]. Following recent
studies of the variability of dental morphology, some
fossil specimens from China have recently been reas-
signed to the genus Meles [5]. A detailed morphologi-
cal description is therefore necessary for isolated teeth
of Arctonyx and Meles, which includes characteriza-
tion of structural variants.
Below is a description of dental remains of badgers
from the Pleistocene deposits of the Lang Trang
karstic cave in northern Vietnam (Thanh Hóa Prov-
ince, Bá Thưc District), collected by the field team of
the Borissiak Paleontological Institute of the Russian
Academy of Sciences (PIN) in March 2020. Thirty-
three specimens of A. collaris have previously been
reported from the Lang Trang cave [6, Table 1], but
without a description or illustrations.
The dental structure terminology, morphotypes,
and measurement technique are according to [5].
Dimensions are indicated in mm. Abbreviations of
measurements: L, maximum length; EL, labial length
(paracone to metacone or paracone to metastyle if
present); IL, lingual length; ETRL, trigonid labial
length (paraconid to hypoflexid); ITRL, trigonid lin-
gual length (paraconid to metaconid); W, maximum
width; TRW, trigonid width; TALW, talonid width;
μ,average value.
The specimens are stored in the collections of PIN,
Moscow. They include two isolated M1 (specimen
PIN, nos. 5792/1, 5792/55) and one М1 on a small
fragment of the dentary (specimen PIN, no. 5792/2).
М1 (specimen PIN, no. 5792/1) has characters
more typical of Arctonyx than Meles [5]. The crown is
rel atively narrow (Fig. 1a ). The labial incis ion b etween
paracone and metacone is weakly developed (mor-
photype A2), the incision posterior to the metaconule
is moderately developed (morphotype C2). No labial
incision is present between the metacone and the
metaconule (morphotype B1), but there is a slight fold
framed by a weakly developed cingulum-like metastyle
(morphotype D2). The lingual incision is absent. The
anterolingual corner of the lingual cingulum projects
anteriorly approximately to the level of the apex of the
paracone (morphotype E4). The posterolabial crest
(between the distal cusp of the protocone and the
metaconule) is absent (morphotype G1), posterolin-
gual crest (between the distal cusp of the protocone
and the hypocone) is weakly developed, incomplete
(morphotype H2). On the talon, there is one relatively
large accessory cusp near the metaconule (morpho-
type F3) and another small accessory cusp near the
hypocone.
Measurements of М1 (specimen PIN, no. 5792/1):
L, 17.0; EL, 13.3; IL, 15.5; W, 13.0. The range of vari-
GENERAL BIOLOGY
a Borissiak Paleontological Institute, Russian Academy
of Sciences, Moscow, 117647 Russia
*e-mail: alopat@paleo.ru
DOKLADY BIOLOGICAL SCIENCES Vol. 495 2020
GREATER HOG BADGER ARCTONYX COLLARIS (MUSTELIDAE, CARNIVORA) 281
ability in М1 measurements for modern representa-
tives of the species [1, Table 3]: L, 14.1–19.1 (μL,
16.7); W, 9.4–12.2 (μW, 11.2). Measurements of the
М1 of A. c. rostratus from the Pleistocene of China [4,
Table 2]: L, 16.0–20.0 (μL, 18.0); W, 10.6–12.9 (μW,
11. 8 ).
Specimen PIN, no. 5792/55 (broken in the
paracone and metacone region) has a narrow talon
with five small accessory cusps (morphotype F2). The
incision posterior to the metaconule is well developed
(morphotype C3). The lingual incision is very slight
(Fig. 1b). The anterolingual corner of the lingual cin-
gulum projects anteriorly significantly less than the
paracone. The posterolabial crest is absent (morpho-
type G1), the posterolingual crest is distinct (morpho-
type H3).
М1 (specimen PIN, no. 5792/2) is long and narrow
(Fig. 2). The paraconid strongly projects anteriorly. A
carnassial notch between the paraconid and protoco-
nid is absent; a short blade is present on the anterior
side of the protoconid. The anterior margin of the
hypoconid is located approximately on the same line
as the apex of the metaconid (morphotype A2),
whereas the apex of the hypoconid is somewhat dis-
placed anteriorly relative to the apex of the mesial
entoconid. The posterior part of the talonid is framed
by the hypoconulid with an extended posterolingual
projection, the distal entoconid and one relatively
large accessory talonid cusp (morphotype B1).
Measurements of М1 (specimen PIN no. 5792/2):
L, 21.2; ETRL, 10.0; ITRL, 12.5; TRW, 6.5; TALW,
7.4. The range of variability in М1 measurements for
modern representatives of the species [1, Table 3]: L,
15.2–20.1 (μL, 18.1 ); TALW, 6.17.7 (μTALW, 6.8).
Same values in A. c. rostratus from various Pleistocene
localities in China [4, Table 2]: L, 16.8–21.0 (μL,
17.9); TALW, 6.5–8.0 (μTALW, 7.0).
Clearly, the L ang Trang form is larger than modern
greater hog badgers in M1 length (specimen PIN,
no. 5792/2) and М1 width (specimen PIN no. 5792/1)
and is only somewhat larger than the known speci-
Fig. 1. Arctonyx collaris rostratus Matthew et Granger,
1923: (a) specimen PIN, no. 5792/1, right M1, occlusal
view, (b): specimen PIN, no. 5792/55, left M
1, occlusal
view; Vietnam, Thanh Hóa Province, Bá Thưc District,
Lang Trang cave; Upper Pleistocene.
0 10 mm
(a)
(b)
Fig. 2. Arctonyx collaris rostratus Matthew et Granger,
1923, specimen PIN, no. 5792/2, right dentary fragment
with M1 in labial (above), occlusal (middle) and lingual
(below) views; Vietnam, Thanh Hóa Province, Bá Thưc
District, Lang Trang cave; Upper Pleistocene.
0 10 mm
282
DOKLADY BIOLOGICAL SCIENCES Vol. 495 2020
LOPATIN
mens of A. c. rostratus. Therefore, based on the size
and structure of the teeth, it can be assigned to the
subspecies A. c. rostratus.
The fossil A. collaris rostratus was initially described
from China as a separate species A. rostratus Matthew
et Granger, 1923, which was considered larger than
A. collaris [7, Figs. 12–14]. This was based on compar-
ison of A. rostratus with specimens of a modern popu-
lation from Central China (Shaanxi), which is now
assigned to a separate species of northern hog badgers,
A. albogularis (Blyth, 1853) [1]. The condylobasal
skull length of A. c. rostratus, however, is 148 mm [7],
which is close to the upper range of A. albogularis
(110–149 mm) and is even somewhat less than in the
modern A. collaris (150–174 mm) [1, 2]. At the same
time A. c. rostratus has relatively large and massive
cheek teeth. Earlier, many findings in the Pleistocene
of Indochina and Thailand were identified as A. col-
laris [6, 8], A. collaris cf. rostratus [9, 10], and A. c. ros-
tratus [11, 12], and it can therefore be assumed that the
geographic distribution of the chronological subspe-
cies A. c. rostratus encompassed, in addition to central,
eastern and southern parts of China [4, 13], also
Southeast Asia.
Arctonyx collaris is characterized by very high
intrapopulational variation in the size and structure of
cheek teeth that is unusual for Carnivora [1, 5, 14].
This may be explained by vermivory, strongly devel-
oped in this species (even compared with other bad-
gers), i.e. feeding on earthworms [1], which some-
times form up to 70–100% of its diet [2]. Vermivory in
mammals often correlates with extreme morphologi-
cal variability of molars, supposedly ref lecting a
reduction in the genetic control on molar develop-
ment in response to a relaxed selective pressure for
exact dental occlusion [1, 15]. Fossil evidence indi-
cates that the Pleistocene populations of A. collaris
generally had equally high morphological and mor-
phometric variability of cheek teeth.
ACKNOWLEDGMENTS
The author thanks E.N. Maschenko, N.V. Serdyuk,
A.A. Lozovsky (PIN), and Le Xuan Dac (Institute of Trop-
ical Ecology of the Joint Russian-Vietnamese Tropical Sci-
entific and Technological Center, Hanoi, Vietnam) for par-
ticipation in collecting of materials in the Lang Trang cave
in March 2020, A.N. Kuznetsov and S.P. Kuznetsova
(Tropical Center) for their assistance in organizing the
expedition, A.M. Kuznetsov and S.V. Bagirov (PIN) for
their help in preparing photographs.
FUNDING
The study was carried out as part of the work of the Joint
Russian–Vietnamese Tropical Scientific and Technological
Center for 2020–2022 (project “The Composition of the
Fauna of Primates (Cercopithecidae, Pongidae) and
Rodents (Rodentia) of the Pleistocene and Holocene of
Vietnam as an Indicator of Environmental Change”).
COMPLIANCE WITH ETHICAL STANDARDS
The study does not contain any research involving ani-
mals or humans.
CONFLICT OF INTEREST
The author declares that he has no conflict of interest.
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Translated by D. Ponomarenko
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“Arctonyx” fossil remains from the Liucheng Gigantopithecus Cave, Guangxi, are redescribed and analysed in details. Detailed tooth character differences between Arctonyx and Meles are analysed. It is shown that materials from the Liucheng Gigantopithecus Cave actually belong to two species of Meles: Meles minor and Meles magnus n. sp. At the same time, a review of Late Pliocene and Early Pleistocene Meles records in China is made. During Late Pliocene, Meles are only represented by M. chiai and one archaic form. During Early Pleistocene, Meles from northern and central part of China are represented by two nearly sympatric species Meles chiai and Meles teihardi. Meles from South China are represented by M. minor and M. magnus n. sp., though the distribution of the two species is still unclear. M. magnus n. sp. is so far only known from the Liucheng Gigantopithecus Cave, whereas M. minor is probably also known from Longgupo, Chongqing in the central part of China besides Liucheng. Great diversity of Meles in Early Pleistocene in China indicates that the genus radiated earlier than previously thought. Phylogenetic analysis suggests M. magnus n. sp. is sister group to living M. leucurus, whereas M. minor and M. chiai are early branches in Meles evolution.
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The extant mammals have evolved highly diversified diets associated with many specialized morphologies. Two rare diets, termitophagy and vermivory, are characterized by unusual morphological and dental adaptations that have evolved independently in several clades. Termitophagy is known to be associated with increases in tooth number, crown simplification, enamel loss, and the appearance of intermolar diastemata. We observed similar modifications at the species level in vermivorous clades, although interestingly the vermivorous mammals lack secondarily derived tools that compensate for the dentition's reduced function. We argue that the parallel dental changes in these specialists are the result of relaxed selection on occlusal functions of the dentition, which allow a parallel cascade of changes to occur independently in each clade. Comparison of the phenotypes of Rhynchomys, a vermivorous rat, and strains of mice whose ectodysplasin (EDA) pathway has been mutated revealed several shared dental features. Our results point to the likely involvement of this genetic pathway in the rapid, parallel morphological specializations in termitophagous and vermivorous species. We show that diets or feeding mechanisms in other mammals that are linked to decreased reliance on complex can lead to similar cascades of change.
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Hog-badgers (mustelid carnivorans classified in the genus Arctonyx) are distributed throughout East and Southeast Asia, including much of China, the eastern Indian Subcontinent, Indochina and the large continental Asian island of Sumatra. Arctonyx is usually regarded as monotypic, comprising the single species A. collaris F. Cuvier, 1825, but taxonomic boundaries in the genus have never been revised on the basis of sizeable series from throughout this geographical range. Based on a review of most available specimens in world museums, we recognize three distinctive species within the genus, based on craniometric analyses, qualitative craniodental features, external comparisons, and geographical and ecological considerations. Arctonyx albogularis (Blyth, 1853) is a shaggy-coated, medium-sized badger widely distributed in temperate Asia, from Tibet and the Himalayan region to eastern and southern China. Arctonyx collaris F. Cuvier, 1825, is an extremely large, shorter-haired badger, distributed throughout Southeast Asia, from eastern India to Myanmar, Thailand, Vietnam, Cambodia and Laos. The world's largest extant badger, A. collaris co-occurs with A. albogularis in eastern India and probably in southern China, and fossil comparisons indicate that its geographical range may have extended into central China in the middle Pleistocene. The disjunctly distributed species Arctonyx hoevenii (Hubrecht, 1891), originally described within the order ‘Edentata’ by a remarkable misunderstanding, is the smallest and darkest member of the genus and is endemic to the Barisan mountain chain of Sumatra. Apart from A. hoevenii, no other Arctonyx occurs on the Sunda Shelf below peninsular Thailand. The natural history of each species of Arctonyx, so far as is known, is briefly reviewed. No claim to original US Government works. Journal compilation © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 353–385.
Article
Thailand has long held a key position in South–East Asia because of its location at the boundary of the Indochinese and Sundaic provinces, the major biogeographical regions of South–East Asia. These provinces are distinct climatically, floristically and faunistically. The present-day limit between them is located at the Kra Isthmus, in peninsular Thailand.Previous studies of the Javanese large mammal fossil faunas and the recent study of fossil large mammal faunas from Thailand strengthen the hypothesis of a “continental” migration route (in contrast with the “insular” hypothesis via Taiwan and the Philippines) during the Late Middle/Late Pleistocene period. Thailand was even part of this migration route. During the glacial periods, the faunal exchanges were favored by the emersion of a huge continental shelf called Sundaland (South–East Asian continental area connected to Borneo and Indonesia islands by land bridges), when the sea level was low. No geological, biogeographical or paleobiogeographical evidence supports the hypothesis of a migration route via Taiwan and the Philippines.Analysis of the extant and Late Middle Pleistocene large mammal faunas (Carnivora, Primates, Proboscidea and Ungulata) points out the antiquity of the Indochinese and Sundaic provinces. This idea is also supported by the fact that mainland faunas already displayed modern character, whereas the Javanese faunas were mainly composed of endemic forms. However, the occurrence of the extinct species, Elephas namadicus found in a Malaysian site (Tambun) indicates that the limit between the two provinces, if not close to the present place, was located more southward than today, possibly in peninsular Malaysia. It is also confirmed by the presence of northern species, such as Ailuropoda melanoleuca baconi and Crocuta crocuta ultima, in Late Middle Pleistocene sites of Thailand. In fact, because of climatic cooling that occurred in the northern hemisphere during the Pleistocene, the northern faunas had to move southward. To our knowledge, the hypothesis of the antiquity of the South–East Asian provinces with a boundary south of the present-day one is mentioned here for the first time.Previous biogeographical studies were based mainly on insular data from the Philippine Archipelago (geological and mainly present-day zoogeographical data) but also from Indonesia and Borneo (paleontological data). However, the recent discovery of Thai large mammal faunas from Late Middle Pleistocene allows to bridge the gap in the fossil record between the northern faunas of South–East Asia and the southern ones. These Thai faunas are characterized mainly by extant forms but some are today absent from the Thai territory (Ailuropoda melanoleuca, Crocuta crocuta, Pongo pygmaeus, Sus cf. barbatus, Rhinoceros cf. unicornis, Cervus eldii).
  • K M Helgen
  • N T Lim
  • L E Helgen
Helgen, K.M., Lim, N.T., and Helgen, L.E., Zool. J. Linn. Soc., 2008, vol. 154, pp. 353-385.
  • H Tong
  • X Wu
  • Z Dong
Tong, H., Wu, X., Dong, Z., et al., Acta Anthropol. Sin., 2018, vol. 27, no. 2, pp. 284-305.