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283
ISSN 0012-4966, Doklady Biological Sciences, 2020, Vol. 495, pp. 283–287. © Pleiades Publishing, Ltd., 2020.
Russian Text © The Author(s), 2020, published in Doklady Rossiiskoi Akademii Nauk. Nauki o Zhizni, 2020, Vol. 495, pp. 581–585.
The Systematic Position of a Large Porcupine
(Hystrix, Hystricidae, Rodentia) from the Pleistocene
of Vietnam (Lang Trang Cave)
Academician A. V. Lopatina,*
Received July 2, 2020; revised July 10, 2020; accepted July 10, 2020
Abstract—The dental remains of the porcupine from the Pleistocene deposits of the Lang Trang cave in
northern Vietnam are described. Previously, this porcupine was identified as Hystrix brachyura Linnaeus,
175 8 o r H. indica Kerr, 1792. Based on the analysis of morphometric data, the systematic position of the form
from Lang Trang and other large Pleistocene porcupines of Vietnam is determined as Hystrix kiangsenensis
Wang, 1931.
Keywords: Hystrix brachyura, H. subcristata, H. indica, H. kiangsenensis, Hystricidae, Rodentia, Old World
porcupines, Pleistocene, Lang Trang cave, Vietnam
DOI: 10.1134/S001249662006006X
At present, the common porcupine species of the
genus Hystrix in Southeast Asia is the Malayan porcu-
pine H. brachyura Linnaeus, 1758. It is represented by
three extant subspecies: H. b. hodgsoni Gray, 1847
(Nepal, eastern India), H. b. subcristata Swinhoe, 1870
(Assam in India, Southern China, Singapore, Myan-
mar, Thailand, Laos, Vietnam, and Cambodia) and
the nominal subspecies (Malacca, Sumatra, and Kali-
mantan) [1]. Middle to Late Pleistocene forms that
occurred within the range of the Stegodon–Ailuropoda
fauna include H. b. punungensis Van Weers, 2003 (fos-
sil subspecies) in Java [2], H. b. subcristata in China (as
H. subcristata, H. cf. subcristata) [3, 4] and Thailand
(as H. hodgsoni subcristata) [5], H. brachyura ssp. in
Vietnam and Sumatra [e.g., 6–8]. Further species
known from the Middle to Late Pleistocene of the
same region include H. kiangsenensis Wang, 1931
(China), H. refossa Gervais, 1852 (= H. magna Pei,
1987, China; H. gigantea Van Weers, 1985, Java), and
H. lagrelli Lönnberg, 1924 (China and Java) [2]. Sys-
tematic identification of isolated teeth of fossil porcu-
pines is to a large extent based on their morphometric
characteristics.
Below is a description of dental remains of Pleisto-
cen e porcupin es from the Lang Trang cave in northern
Vietnam (Thanh Hóa Province, Bá Thước District),
collected by the field team of the Borissiak Paleonto-
logical Institute of the Russian Academy of Sciences
(PIN) in March 2020. Previously, 358 specimens of
Hystrix (Acanthion) brachyura were reported from the
Lang Trang cave without description or figures [6,
Table 1]; the same fossil material is apparently men-
tioned under the name Hystrix subcristata [6, Table 2].
Measurements of porcupine teeth from the Lang
Trang cave were published later [2, 7].
The dental structure terminology is according to
[9]; the tooth crown wear stages (classes) are accord-
ing to [10]. Dimensions are indicated in mm.
Measurement abbreviations: LD, longitudinal diame-
ter; TD, transverse diameter; L, maximum length;
OL, occlusal length; OW, occlusal width; IH, lingual
enamel height of the tooth crown; EH, labial enamel
height of the tooth crown; HH, hypostria (hypostriid)
height; HI, paraf lexus (anteroflexid) height; HII,
anterior mesoflexus (anterior mesof lexid) height;
HIII, posterior mesof lexus (posterior mesof lexid)
height; HIV, postflexus (posteroflexid) height;
HV, labial mesoflexid height; n, number of speci-
mens; μ, average value.
The specimens are housed in the collections of
PIN, Moscow. Material includes incisor fragments
and isolated hypsodont premolars.
Measurements of I1 (specimen PIN, no. 5792/99):
LD, 8.5; TD, 6.5. As many other specimens from the
Lang Trang cave, this specimen has gnawing traces left
on it by incisors of other porcupines as part of osteo-
phagic behavior typical to these rodents (Fig. 1).
DP4 (specimen PIN, no. 5792/97) moderately
worn, wear stage D5 (two laterally closed fossettes,
three open folds). The parafossette is linked with the
anterior mesoflexus (the protoloph is interrupted),
GENERAL BIOLOGY
a Borissiak Paleontological Institute, Russian Academy
of Sciences, Moscow, 117647 Russia
*e-mail: alopat@paleo.ru
284
DOKLADY BIOLOGICAL SCIENCES Vol. 495 2020
LOPATIN
isolated from the hypoflexus and closed anteriorly, but
the dentine fields of the anteroloph and the labial part
of the protoloph are separated by enamel walls (Fig. 2).
The anterior mesoflexus is connected with the hypo-
flexus. The posterior mesoflexus is open labially and
confluent with the lingual mesofossette which is
closed distally. The dentine fields of the hypocone and
the posteroloph are connected. The protofossette is
closed, the dentine fields of the metaloph and the
posteroloph are connected by a very narrow bridge.
The lingual root is large, whereas the two labial ones
are relatively small, and the posterior labial one is a
double root. Measurements of DP4 (specimen PIN,
no. 5792/97): OL, 8.6; OW, 5.6; IH, 11.0; HH, 2.3;
HII, 0.4; HIII, 0.2.
P4 (specimen PIN, no. 5792/96) is slightly worn,
wear stage B1 (all reentrant folds are open, no closed
enamel islands). The labialmost part of the anteroloph
is unworn. The paraflexus is linked with the hypo-
flexus and the anterior mesoflexus, the protoloph is
incomplete. The anterior mesoflexus is connected
with the lingual mesofossette and the posterior meso-
flexus. The dentine fields of the hypocone and the
posteroloph are confluent. The metaloph is incom-
plete, the postfossette has two narrow links to the pos-
terior mesoflexus. There is a contact area for М1 on
the posterior side. Roots are absent. Measurements of
P4 (specimen PIN, no. 5792/96): L, 8.7; OL, 8.5; OW,
4.7; IH, 16.5; HH, 10.2; HI, 1.4; HII, 4.3; HIII, 1.5.
Measurements of I1: LD, 6.5; TD, 5.8 (specimen
PIN, no. 5792/98); LD, 6.8; TD, 5.8 (specimen PIN,
no. 5792/127).
The unworn (stage O1) longitudinally elongate
occlusal surface of P4 (specimen PIN, no. 5792/95)
consists of fully enamel-covered cusps (protoconid,
metaconid, central conid, mesostylid, hypoconid,
entoconid, central eminences of the metalophid and
hypolophid) divided by folds, and the anterolophid
and posterolophid crests that are shallowly dissected
by superficial grooves. All the folds are open, the
hypoflexid is broad, the hypostriid very deep. Visible
from the side of the broken roots are the bottoms of
four closed folds of the large elongated hypofossettid
and the smaller posterofossettid, anterior, and poste-
rior mesofossettids. The enamel of the lateral sides of
the tooth is distinctly wrinkled, especially at the base
of the crown. Measurements of P4 (specimen PIN,
no. 5792/95): L, 10.1; OL, 8.8; OW, 6.0; EH > 15.0;
HH > 14.0; HI, 1.4; HII, 0.8; HIII, 2.9; HIV, 0.5;
HV, 1.1.
By analyzing a large sample, van Weers [2] deter-
mined that the porcupine from the Lang Trang cave
has much greater premolar length [2, Fig. 3] than all
fossil and modern H. brachyura sensu lato, and it is
even somewhat greater than the modern H. indica
(Table 1). Differences in molar length are somewhat
less pronounced [2, Fig. 4]. Finds from other Late
Pleistocene cave localities also indicate that the fossil
porcupine from Vietnam was relatively large-sized
[2, 7, 8].
Based on the size of the teeth, van Weers [2] deter-
mined that porcupines from the Pleistocene of Viet-
nam do not belong to H. brachyura, but can be
assigned to H. indica. Van Weers noted, however, that
tooth size also shows similarity with H. kiangsenensis
from the Pleistocene of China, but the lack of infor-
mation on skull morphology of the Vietnamese form
precludes a comparison with the fossil Chinese spe-
cies.
In my opinion, the fossil porcupine from the Lang
Trang cave should not be assigned to H. indica,
because the two species have a different ratio of pre-
molar to molar length. In the Vietnamese form, the
ratio μOL P4/M1–2 equals 1.224, μOL P4/M1–2 equals
1.119, whereas in the Indian porcupine the two ratios
are 1.1 and 1.083 respectively (data from Table 1). The
ratio of maximum OL values is even more striking: in
the Vietnamese form, P4/M1–2 = 1.298, P4/M1–2 =
1.13, whereas in H. indica P4/M1–2 = 1.065, P4/M1–2 =
1.074. The ratio of premolar length to the length of the
first or second molar is therefore considerably greater
in the Vietnamese porcupine than in H. indica.
The assignment of the Vietnamese form to
H. brachyura (including the fossil populations of Java
and Sumatra) is incorrect due to the large difference in
tooth size noted by van Weers. Also, in H. brachyura
sensu lato, μOL P4/M1–2 = 1.136 and μOL P4/M1–2 =
1.113, and the ratio of maximum values OL P4/M1–2 =
1.0 63 and P4/M1–2 = 1.022, i.e., also much less than in
the Vietnamese form.
Noting the relatively large size, relatively high skull
and long nasals, van Weers and Zheng [11] assigned
Middle Pleistocene porcupines from China to the spe-
cies H. kiangsenensis within the subgenus Hystrix,
Fig. 1. Hystrix kiangsenensis Wang, 1931: specimen PIN,
no. 5792/99, fragment of left I1 (above), labial view; spec-
imen PIN, no. 5792/98, fragment of right I1 (below), labial
view; Vietnam, Thanh Hóa Province, Bá Thước District,
Lang Trang Cave; Upper Pleistocene.
0 10 mm
DOKLADY BIOLOGICAL SCIENCES Vol. 495 2020
THE SYSTEMATIC POSITION OF A LARGE PORCUPINE 285
Fig. 2. Hystrix kiangsenensis Wang, 1931: specimen PIN, no. 5792/97, right DP4 (left row); specimen PIN, no. 5792/96, right P4
(middle row); specimen PIN, no. 5792/95, right P4 (right row); occlusal view (above), labial view (middle) and lingual view
(below); Vietnam, Thanh Hóa Province, Bá Thước District, Lang Trang Cave; Upper Pleistocene.
05 mm
while previously they were commonly treated as
H. cf. subcristata [12, 13]. Van Weers later [2] assigned
to H. kiangsenensis various Late Pleistocene porcupine
dental and cranial remains from China [3], retaining
in H. brachyura only the relatively small form from the
Early Pleistocene locality of Longgupo (southern part
of Central China) [14]. He did not exclude the possi-
bility of H. brachyura occurring in other Chinese
localities, including those that contain H. kiangsenen-
sis [2].
This view is rejected by Tong [4], who argues that
these fossil Chinese forms belong to the recent H. sub-
cristata as a separate species for which this author sug-
gests an affinity with Hystrix, rather than Acanthion,
and is characterized, among other things, by species
characters of H. kiangsenensis according to van Weers,
including long nasals and a high skull. In addition,
Tong indicates the large elongated upper premolar as
one of the diagnostic characters for this form [4],
while his morphometric data [3, Table 3] show that the
lower premolar is also relatively large and elongated
(μOL P4/M1–2 = 1.176, P4/M1 = 1.329, P4/M2 = 1.296,
maximum values OL P4/M1–2 = 1.188, P4/M1 = 1.263,
P4/M2 = 1.333).
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DOKLADY BIOLOGICAL SCIENCES Vol. 495 2020
LOPATIN
It should be noted here that extant H. b. subcristata,
being a relatively large representative of H. brachyura
sensu lato [1], is nonetheless considerably smaller than
H. kiangsenensis in maximum and average cheek teeth
length [2], whereas molecular genetic evidence does
not support species level for H. b. subcristata [15].
In conclusion, the totality of morphometric evi-
dence (the size of cheek teeth and the relative length of
the premolars) indicate that the porcupine from the
Lang Trang cave should be treated as Hystrix kiangsen-
ensis Wang, 1931 (sensu van Weers). This identifica-
tion should also be extended to other relatively large
Pleistocene porcupines of Southeast Asia, namely
“H. brachyura” from Vietnam [7, 8] and “H. cf. indica”
(= “H. hodgsoni subcristata”) from Thailand [5].
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.
REFERENCES
1. Van Weers, D.J., Beaufortia, 1979, vol. 29, no. 356,
pp. 215–272.
2. Van Weers, D.J., Contrib. Zool., 2005, vol. 74, no. 3/4,
pp. 301–312.
3. Tong, H.W., Vertebr. PalAsiat., 2005, vol. 43, no. 2,
pp. 135–150.
4. Tong, H., Quat. Int., 2008, vol. 179, pp. 126–134.
5. Filoux, A., Wattanapituksakul, A., Lespes, C., and
Thongcharoenchaikit, C., Geobios, 2015, vol. 48,
pp. 341–349.
Table 1. Comparison of tooth measurements for porcupines of the genus Hystrix from Vietnam (the caves of Lang Trang,
Thẩm Khuyên, Thẩm Ồm, and Đười Ươi, Upper Pleistocene) and other countries and regions of Asia
Taxon, locality Р4M1–2 P4M1–2
OL μOL (n) OL μOL (n) OL μOL (n) OL μOL (n)
Lang Trang [2] 8.3–10.9 9.3 (21) 6.7–8.4 7.6 (20) 8.6–10.4 9.4 (11) 7.5–9.2 8.4 (23)
Lang Trang [7] ––––8.0–10.1– (10)7.5–9.2– (23)
Lang Trang (our data) 8.5–––8.8–––
Thẩm Khuyên [2] –9.1 (6)–8.0 (11)––––
Thẩm Ồm [2] –8.7 (9)–7.6 (13)––––
Ma Ươi [7] ––––8.25– (1)8.07–8.18.09 (2)
Đười Ươi [8] ––––7.2–9.1– (22)8.4–11.9– (30)
Tham Prakai Phet, Pleistocene
of Thailand [5] – – – 8.2 (8) – 10.5 (3) – 8.7 (5)
H. kiangsenensis, Pleistocene
of China [2] 7.0–10.0 8.8 (25) 6.3–10.2 7.9 (53) 8.8–11.3 9.8 (22) 6.3–9.7 8.4 (20)
H. kiangsenensis, Tianyuan,
China, Upper Pleistocene [3] 8.0–9.5 8.7 (5) 6.0–8.0 7.4 (5) 8.5–12.0 10.5 (3) 7.0–9.5 – (13)
H. brachyura s.l., extant [2] 6.5–8.5 7.5 (32) 4.7–8.0 6.6 (91) 6.4–9.1 7.9 (30) 4.6–8.9 7.1 (87)
H. brachyura, Pleistocene
of Sumatra [2] 6.9–8.9 7.9 (87) 6.0–8.6 6.8 (123) 6.9–9.9 8.4 (69) 5.8–8.8 7.6 (107)
H. brachyura punungensis,
Pleistocene of Java [2] 6.2–8.7 7.6 (52) 5.2–8.0 6.7 (74) 7.1–10.1 8.6 (55) 5.5–8.7 7.3 (72)
H. indica, extant [2] 7.5–9.8 8.8 (20) 6.3–9.2 8.0 (53) 7.4–10.2 9.1 (23) 6.9–9.5 8.4 (55)
DOKLADY BIOLOGICAL SCIENCES Vol. 495 2020
THE SYSTEMATIC POSITION OF A LARGE PORCUPINE 287
6. Long, V.T., De Vos, J., and Ciochon, R.L., Bull. Indo-
Paci f. Prehist. Assoc., 1996, vol. 14, pp. 101–109.
7. Bacon, A.-M., Demeter, F., Roussé, S., et al., Palaeo-
geogr. Palaeoclimatol. Palaeoecol., 2006, vol. 230,
pp. 280–298.
8. Bacon, A.-M., Demeter, F., Duringer, P., et al., Quat.
Sci. Rev., 2008, vol. 27, pp. 1627–1654.
9. Lopatin, A.V., Tesakov, A.S., and Titov, V.V., Rus. J.
Theriol., 2003, vol. 2, no. 1, pp. 26–32.
10. Van Weers, D.J., Bijdrag. Dierk., 1990, vol. 60, no. 2,
pp. 121–134.
11. Van Weers, D.J. and Zheng, S., Beaufortia, 1998,
vol. 48, no. 4, pp. 47–69.
12. Young, C.C., Palaeontol. Sin. Ser. C, 1934, vol. 8, no. 3,
pp. 30–41.
13. Colbert, E.H. and Hooijer, D.A., Bull. Am. Mus. Nat.
Hist., 1953, vol. 102, pp. 1–134.
14. van Weers, D.J., Beaufortia, 2003, vol. 53, no. 3,
pp. 61–65.
15. Rovie-Ryan, J.J., Anwarali Khan, F.A., Zainuddin, Z.Z.,
et al., J. Sustain. Sci. Manag., 2017, vol. 12, no. 1,
pp. 1–11.
Translated by D. Ponomarenko
