Hystrix zhengi n. so., a brachyodont Porcupine (Rodentia) from early Nihewanian stage, early Pleistocene of China

8 Pages·1999·1.5 MB·English

by  Van Weeks, D J

Beaufortia
INSTITUTE FOR SYSTEMATICS AND POPULATIONBIOLOGY
(ZOOLOGICAL MUSEUM) UNIVERSITYOFAMSTERDAM
Vol. 49, no. 7 December 17, 1999
Hystrixzhengin. so., a brachyodontPorcupine(Rodentia) from
early Nihewanianstage, early Pleistocene ofChina
D.J. vanWeeks & ZhangZhaoqun
*InstituteforSystematicsandPopulation Biology (Zoological Museum), UniversityofAmsterdam,
P.O. Box 94766, 1090GTAmsterdam, TheNetherlands
**InstituteofVertebratePaleontology andPaleoanthropology, AcademiaSinica,
P.O. Box643,Beijing 100044,PR. China
Keywords: Hystrix, China, Nihewanian, Early Pleistocene, Late Pliocene, H. zhengi, H. magna, H.
kiangsenensis
Abstract
A lowerpremolar fromthe LonggupoCave, Sichuan,China and three lowerpremolars fromtheLiuchengGigantopithecus
Cave, Guangxi,are describedas Hystrixzhengin.sp. The localities Longgupoand Liucheng areofEarly Nihewanianage,
which correlates with the Late Pliocene (Villanyian) ofEurope. The specimens are considered torepresenta late occur-
renceofa brachyodontspecies,agreeingwith the replacementofbrachyodontHystrixspeciesbyhypsodontonesin Europe
duringthe Late Pliocene.
Chineseabstract
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55
INTRODUCTION SPECIMENSSTUDIED
The Longgupo Cave, also known as the "Wu- IVPPV9669, leftp4, Longgupo, fissure zone 6.
shan Hominid Site", in Sichuan Province, IVPPVI 1549.1, left p4, Liucheng Gigantopithecus
China, yielded one brachyodont lower premolar Cave.
ofHystrix. Itwas found during the excavations of IVPPVI 1549.2, left p4, Liucheng Gigantopithecus
1985-1988and was described by Zheng (1993) Cave.
under the name Hystrix magnaPei, 1987. The IVPP VI1549.3, right p4, Liucheng Gigantopithe-
type material ofH. magnawas revised by van cusCave.
Weers & Zheng (1998) who designated the only IVPP VI 1550.1, H. magna, left p4, Liucheng
syntype thatwas left as the lectotype for H. mag- Gigantopithecus Cave.
na. This tooth, however, is hypsodont. The clear- IVPP VI 1550.2, H. magna, right P4, Liucheng
ly lower crowned p4 from Longgupo is outside Gigantopithecus Cave.
the range ofvariation of the tooth height of H. IVPP VI 1550.3, H. magna, left p4, Liucheng
magna.Therefore the specific designation of that Gigantopithecus Cave.
tooth willbereconsidered. IVPP V11551.1, H. kiangsenensis, right p4, Liu-
While searching in vain for syntypes ofH. cheng Gigantopithecus Cave.
magna in the collections of the IVPP, three bra- IVPP VI 1551.2, H. kiangsenensis, left p4, Liu-
chyodont lowerpremolars of aporcupine species cheng Gigantopithecus Cave.
from the Liucheng Gigantopithecus Cave were
found. In this paper the four molars mentioned TAXONOMY
above will be compared with the molars of the
species H. magnaand H. kiangsenensis Wang, 1931 OrderRodentiaBowdich, 1821
from the Gigantopithecus Cave. Family Hystricidae Burnett, 1830
GenusHystrix Linnaeus, 1758
MATERIALSAND METHODS Subgenus Hystrix Linnaeus, 1758
This subgenus encloses the larger sized Hystrix
MEASUREMENTS species, hypsodont as well as brachyodont ones,
which are more specialised as for their larger
Thelargest breadth(Br) and length (Lth), as well length ofthe nasals and the larger height of the
as three differentmeasurements ofthe height of skull.
the teeth havebeen taken with vernier callipers.
Thetotal height (Ht) is the height withthe roots. Hystrix zhengi n. sp.
Anothermeasurement taken is the smallest (Fig 1A-I)
height of the tooth without the roots ("crown
height" = C.Ht). This measurement has the Hystrix magna Pei, 1987: 115(specimens nowreg-
advantage thatit canalways be takeneven ifthe istered as VI 1549.1-3); Zheng, 1993: 24, Fig.
enamel/dentineborder is not marked. The 57a.
enamel height (E.Ht) is the largest heigt of the Hystrix sp.; vanWeers & Zheng, 1998:55.
enamel crown, in lower teeth at the buccal, in
upper teeth at the lingual side. The relative Holotype: left lowerpremolar kept inthe IVPP in
height of the teeth is expressed as the ratio of Beijing undernr. V9669, Figs. 1A-C.
E.Ht/Lth. Therecognition ofthe wear classes is Derivatio nominis: named after Dr. Zheng
after van Weers (1990). Upper cheek teeth are Shaohua, acknowledging his contribution to the
indicatedin uppercase,lower ones in lowercase. Quaternary mammology.
First andsecond lowermolars are together treat- Type locality: Site ofLonggupo, fissure zone 6,
edas ml/2. Wushan County, SichuanProvince, China.
Hypodigm: V9669, holotype, left p4 (Fig. 1A-C);
VI 1549.1 (Fig. 1D-E) and VI 1549.2(Fig. 1F-G),
leftp4's, andVI 1549.3, right p4(Fig. 1H-I).
56
Fig. 1.A,occlusal,B, buccal and C, lingualview oftheleftp4,holotypeofH. zhengin. sp.nr.IVPPV9669 fromLonggupo.
D,occlusal and E, buccalview ofleft p4nr.IVPPV11549.1;F, occlusal and G, buccal view ofleftp4nr. IVPPV11549.2;
H,occlusal and I, buccalview ofrightp4IVPPV11549.3 ofH. zhengin.sp.fromtheLiuchengGigantopithecusCave.
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Table 1.Measurements ofthe type(V9669) andthreep4ofH. zhengi,twop4and oneP4ofH. magnaand twop4ofH.
kiangsenensis inmm. Heightmeasurementsofteeth with damagedorlackingroots areindicated with*.
IVPPnr. Locality Species Th Br Lth Ht C.Ht E.Ht CI Ht/Lth
V9669 Longgupo H. zhengi P4 9.0 12.2 17.1 10.7 11.0 PI 0.90
V11549.1 Liucheng H. zhengi p4 8.8 11.0 12.5* 9.9 11.0 Q. 1.00
VI1549.2 Liucheng H. zhengi p4 8.4 12.0 13.2* 11.8 11.8 T1 0.98
VI1549.3 Liucheng H. zhengi p4 9.8 11.5 11.0* 8.6 10.9 Q, 0.95
V11550.1 Liucheng H. magna p4 7.7 10.6 15.3 13.6 13.0 S4 1.23
VI1550.2 Liucheng H. magna P4 9.1 10.5 18.8 19.1 G1 1.82
VI1550.3 Liucheng H. magna p4 8.1 10.2 18.0 16.2 16.9 R/S 1.66
VI1551.1 Liucheng H. kiangsenensis p4 6.6 8.4 14.8* 14.8 PI 1.76
VI1551.2 Liucheng H.kiangsenensis p4 6.7 8.8 16.1 11.4 15.0 PI 1.70
Age: Early Nihewanian, Chinese Early Pleisto- species. That tablealso shows that H. zhengi and
cene,which agreeswith the European Late Plio- H. magna are larger than H. kiangsenensis. The
cene. buccal views ofH. zhengi (Fig. 1) and of H. magna
Diagnosis: H. zhengi is a very large porcupine with and H. kiangsenensis (Fig. 2) illustrate the differ-
brachyodont cheek teeth. ence in height between H. zhengi and the latter
Differential diagnosis: H. zhengi differs from H. two species. The occlusal views in the figures 1
kiangsenensis by its larger size and brachyodont and 2 showthe stage ofwear, the class ofwear is
molars. H. zhengi has about the same size as H. given inTable 1.
magna, but the latter species is clearly hypsodont
with arelative average crown height (E.Ht/Lth) DISCUSSION
ofthe p4of 1.9. H. zhengi differs from theMiddle
Pleistocene H. gigantea van Weers, 1985 ofJava Pei (1987) mentionedmore than 200 isolated
by its brachyodonty. The size ofthe p4 of H. porcupine cheekteeth from the Liucheng Gigan-
zhengi is about the same as in H. primigenia Wag- topithecus Cave. Most oftheseare allocatedto the
ner, 1848but it is less brachyodont. It has arela- hypsodont species H. magnaand .H. kiangsenensis
tive crown height of an average of0.96 against (van Weers & Zheng, 1998). Pei (1987) allocated
about0.7 in H.primigenia. only 14 specimens from that large collection to
H. magnaand considered themas relatively low-
DESCRIPTION crownedand intermediate in height between the
brachyodont Atherurus and hypsodont Hystrix
Table 1 gives the measurements ofthe holotype teeth. The only left one of these syntypes of H.
of H. zhengi from Longgupo and ofthe three p4 magna is a hypsodont Ml or M2, original num-
from the Liucheng Gigantopithecus Cave assigned ber V5036.12, nowkept inthe IVPPas lectotype
to thatspecies. The measurements oftwo p4and under number VI 1549.4. Van Weers & Zheng
one P4 of H. magna and of two p4 of H. kiangse- (1998) considered that Pei's type series of H.
nensis from the Liucheng Gigantopithecus Cave are magna could have been the lowest specimens in
given for comparison. The height ofthe enamel the rangeof variation ofthe specimens allocated
crown ofthe holotype is dependent on the place by themto H. magna. Itmustbe testedifthis sup-
where it is measured. That height is buccally at position is valid for the low-crowned teeth ofH.
midlength 10.2 mm, buccal-posteriorly 11.0mm zhengi. Theonly sample available forcomparison
and anteriorly 11.4mm. At the occlusal surface consists of the mixture ofp4 of H. magnaandH.
the breadth of the tooth is 7.8 mm and the kiangsenensis (van Weers & Zheng, Table 2). These
length 10.6 mm. show arelative height (Ht/Lth) for thep4 of 1.6-
Table 1 shows the differences in the absolute 2.4, mean 1.9 (n = 15), so these are all clearly
and relative tooth height between the three hypsodont and significantly different from H.
58
Fig. 2.A and C, occlusal, B and D, buccal views ofleftp4nr. IVPPV11550.1 and V11550.3;E, occlusal andF, buccal
view ofright P4nr.IVPP V11550.2ofH. magnafromtheLiuchengGigantopithecus Cave. G, occlusal and H,buccal view of
the rightp4nr.IVPPV11551.1;I, occlusalandJ,buccal view ofthe leftp4nr.IVPPV11551.2 ofH.kiangsenensis fromthe
LiuchengGigantopithecus Cave.
59
Fig. 3.Diagram oflength and height of 13lowerpremolars ofthe Mio-Pliocene H.primigeniafromEurope,the holotypeof
H. zhengin.sp.fromLonggupo and threespecimens ofthatspecies fromtheLiuchengGigantopithecusCave.
zhengi. In this light Pei's (1987) concept of H. dont H. sivalensis Lydekker, 1878 from the Late
magna can be better explained by the supposition Miocene of the Siwaliks. This is done indirectly
that his type series consisted of a mixture of H. via H. primigenia. Only from the m1 ofthe holo-
zhengi andH. magna. type mandible of H. sivalensis,, the hypodigm of
The supposition that the small height of the that species, the height and length couldbe mea-
teeth of H. zhengi is dueto the advanced stage of sured. That tooth differs neither in length nor in
wear does not apply because three of the four height from H. primigenia (Fig. 4), so there is no
teeth belong to the rather youngwear classes P basis to suppose a closerrelationship ofH. zhengi
and Ç) (van Weers, 1990). withH. sivalensis thanwithH. primigenia.
H. zhengi resembles the Mio-PlioceneEuro- Theage of the Longgupo fauna is compara-
pean H. primigenia (MN12-MN16) best. Figure 3 ble with the Dachaian fauna ofNorth China,
shows that thelength ofthe fourteethof H. zhen- Late Pliocene, MN17, (Zheng, 1993and Huang
gi does not differ from asample ofthirteenspeci- Wanpo et al., 1995). Theage of the Longgupo
mens of H. primigenia (Rook & van Weers, in and Liucheng faunas are similar as shown by the
prep.). The mean height of the new Chinese co-existence of many genera and species (Pei
species however, differs signiflcandy from that of 1987, HanDefen 1987and Zheng 1993).
the European one. Thehypodigm of H. zhengi is very small. This
H. zhengi is also compared with the brachyo- material probably represents a relic of a Late
60
Fig. 4.Diagram oflengthand height of18first and second lowermolarsofthe Mio-Pliocene H.primigeniafromEuropeand
the firstlowermolarofthe holotype mandibleofH. sivalensisfromtheSiwaliks.
Miocene brachyodont species. The last occur- and Mr Wil denHartog of the same institution
rence of H.primigenia in Europe is inthePliocene mountedthe plates. The exchange of ideas with
of Perpignan, France (Ruscinian, MN15). It Dr John de Vos (National Museum of Natural
seems that the brachyodont Hystrix species of History, Leiden, TheNetherlands) was very use-
Asia and Europe were both replaced by hyp- ful. The financial support ofthe Netherlands
sodont species during the LatePliocene. Organisation for Scientific Research (NWO),
grantR 87-280, is greatly acknowledged.
ACKNOWLEDGEMENTS
REFERENCES
The hospitality ofDr Qiu Zhuding, the director
HAN DEFEN, 1987. Artiodactyla fossils from Liucheng
of the IVPP in Beijing, during my visit is highly Gigantopithecus Cavein Guangxi. Mem. ofInst, ofVert.
appreciated. I am very thankful to Dr Zheng Palaeont.and Palaeoanthr., Acad. Sinica, 18: 135-200
Shaohuafor entrusting to me his collectionsand (Chinese)(Englishsummery: 201-205).
for his kind co-operation. I appreciated the help HUANG WANPO, R. CIOCHON, GU YUM1N, R.
of Dr Wu Wenyu during my stay. Mr Zhang Jie LARICK, FANG QIREN, H. SCHWARCZ, C.
YONGE,J. DE VOS& W.RINCK, 1995. Early Homo
made the photographs. A critical reading by Dr
and associated artefacts from Asia. Nature 378: 275-
Hans de Bruijn (Institute for Earth Science,
278.
Utrecht, The Netherlands) improved the text, PEI WENZHONG, 1987. Carnívora, Proboscidea and
61
Rodentiafrom Liucheng GigantopithecusCave andother WEERS, D.J. VAN & ZHENG SHAOHUA, 1998.
caves in Guangxi. Mem. Inst. Vert. Palaeont. Palaeo- Biométrieanalysis and taxonomie allocation of
anthr., Acad. Sinica, 18: 1-94 (Chinese) (English Pleistocene Hystrix specimens (Rodentia,Porcupines)
abstract:95-115). fromChina. Beaufortia 48(4): 47-69.
WEERS,D.J. VAN, 1985. Hystrix gigantea,anewfossil por- ZHENG SHAOHUA, 1993.Quaternaryrodents ofSichu-
cupine species from Java(Rodentia: Hystricidae). an-Guizhou area, China. Academia Sinica, Beijing: 1-
Senckenbergianalethaea,66 (1-2): 111-119. 230(Chinese)(English summary: 231-270).
WEERS, D.J. VAN, 1990. Dimensionsand occlusal pat-
terns inmolarsofHystrixbrachyura.
Linnaeus, 1758 (Mammalia,Rodentia)in asystem ofwear
categories.Bijdr. Dierk.,60 (2): 121-134. Received: May 3, 1999
62