from the Late Oligocene of north-central Anatolia (Turkey)

12 Pages·2008·0.49 MB·English

Zoological Journal of the Linnean Society, 2008, 152, 581–592. With 4 figures
A giant rhinocerotoid (Mammalia, Perissodactyla) from
the Late Oligocene of north-central Anatolia (Turkey)
PIERRE-OLIVIER ANTOINE1*, LEVENT KARADENIZLI2, GERÇEK SARAÇ3 and
SEVKET SEN4
1Laboratoire des Mécanismes de Transfert en Géologie, Université de Toulouse, CNRS, IRD, OMP, 14
Avenue Édouard Belin, F-31400 Toulouse, France
2Geology Department, Maden ve Tektik Arama Genel Müdürlügü, 06520 Ankara, Turkey
3Natural History Museum, Maden ve Tektik Arama Genel Müdürlügü, 06520 Ankara, Turkey
4Laboratoire de Paléontologie, UMR 5143 CNRS, Muséum National d’Histoire Naturelle, 8 rue
Buffon, F-75005 Paris Cedex 5, France
Received 3 March 2006; accepted for publication 4 June 2007
A giant rhinocerotoid is described for the first time south of the Black Sea, in Turkey. The single specimen, a
fragmentary radius referred to Paraceratherium sp., originates from conglomerates nearby at Gözükizilli, in the
Çankiri–Çorum Tertiary basin. These layers correspond to the Lower member of the Kizilirmak Formation. The
samelocality(Gözükizilli-2)yieldsalsothesmallrhinocerotidProtaceratheriumsp.,cf.P.albigense(Roman,1912).
Three other mammal localities (Gözükizilli-1, in the Lower Member of the formation, with several rodent species;
Tepe 641 and Kizilirmak, in the Upper Member, with a diversified micro- and macro-mammal fauna) allow us to
refer the Kizilirmak Formation as a whole to the Late Oligocene.All the observed taxa have strongAsian and/or
European affinities, which precludes any geographical insulation for this part of Anatolia during the Late
Oligocene. ©2008TheLinneanSocietyofLondon,ZoologicalJournaloftheLinneanSociety,2008,152,581–592.
ADDITIONAL KEYWORDS: indricothere – palaeogeography – Paraceratherium – vertebrate palaeontology.
INTRODUCTION This article deals with the discovery of a single
specimen–abrokenanddamagedradius–unearthed
The indricotheres are extinct giant rhinocerotoids
in June 2002 near the village of Gözükizilli, c.20km
known in the Eocene of western North America and
south of Kizilirmak (Fig.1), during part of a joint
from the Middle Eocene to the Late Oligocene in
project by the Muséum National d’Histoire Naturelle
Eurasia (Russell & Zhai, 1987; Lucas & Sobus, 1989;
in Paris and the Maden Tektik ve Arama in Ankara
Welcomme etal., 2001; Antoine etal., 2004). The
(Karadenizlietal.,2003).Thisradius,unquestionably
hornlessrhinos,whichmanyauthorsconsidertohave
referable to an indricothere (on the basis of size and
been the largest land mammals that have existed,
morphologicalfeatures),isthefirstevidenceofagiant
were widespread during the Oligocene in Asia (Paki-
rhinocerotoid in Asia Minor. It was found associated
stan, China, Mongolia, Russia, Kazakhstan; Forster-
withamagnumofasmallrhinocerotidingreyfluvial
Cooper, 1911, 1913a, b, 1915, 1924, 1934; Borissiak,
conglomerates (Fig.1: Gözükizilli-2 locality; Fig.2)
1915,1923;Pavlova,1922;TeilharddeChardin,1926;
from the Lower Member (Mbr) of the Kizilirmak
Granger & Gregory, 1936; Gromova, 1959; Lucas &
Formation (Fm.). These conglomerates overlie grey
Sobus, 1989; Antoine etal., 2004). In addition, a few
lacustrine clays (Fig.1: Gözükizilli-1 locality), which
specimens have been mentioned in recent decades
have yielded a diversified micromammal fauna,
fromEasternEurope(Yugoslavia,Bulgaria,Romania;
notablyLateOligocenerodents(Ünay,Bruijn&Saraç,
Nikolov & Heissig, 1985; Lucas & Sobus, 1989;
2003).Inaddition,therecentdiscoveryoftwomammal
Spassov, 1989; Codrea, 2000) and the Caucasus
localities(KizilirmakandTepe641;Fig.1)intheupper
(Georgia; Gabunia, 1955, 1964, 1966).
memberoftheKizilirmakFm.shedsanewlightonthe
age of this formation as a whole (Karadenizli etal.,
*Corresponding author. E-mail: [email protected] 2003).
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592 581
582 P.-O. ANTOINE ETAL.
Figure1. Location map of Late Oligocene mammal localities and sections from the Kizilirmak Fm. (Çankiri-Çorum
Basin, north central Anatolia, Turkey). A, map of Turkey; B, location of Kizilirmak, Gözükizilli and other cities; C,
geological sketch map of the Gözükizilli area. GK-1, Gözükizilli-1 (Lower Mbr); GK-2, Gözükizilli-2 (Lower Mbr);
Kizilirmak (Upper Mbr); Tepe 641 (Upper Mbr). In B and C, the coordinates are UTM.
SYSTEMATICS theriinae‘haspriorityoverOsborn’s(1923:13)names
BaluchitheriinaeandParaceratheriinae(imprintdate
The use of the taxon Rhinocerotoidea Gray, 1821
on Borissiak, 1923 is March; on Osborn, 1923, it is
follows Prothero, Guérin & Manning (1989) rather
May)’, according to Lucas & Sobus (1989: 362).
thanMcKenna&Bell(1997):itisthecladeformedby
rhinocerotids,hyracodontidsandamynodontids.Even
ORDER PERISSODACTYLA OWEN, 1848
though the genus Indricotherium Borissiak, 1915 is
SUPERFAMILY RHINOCEROTOIDEA GRAY, 1821
considered here as a junior synonym of Paracerath-
FAMILY RHINOCEROTIDAE GRAY, 1821
eriumForster-Cooper,1911,thesubfamilynameIndri-
GENUS PROTACERATHERIUM ABEL, 1910
cotheriinae Borissiak, 1923 is used (1) because ‘when
the name of a type genus of a nominal family group PROTACERATHERIUM SP., CF. PROTACERATHERIUM
taxonisconsideredtobeajuniorsynonymofthename ALBIGENSE (ROMAN, 1912)
of another nominal genus, the family group name is FIGURE2A–C
not to be replaced on that account alone’(ICZN 1999: Material available: GK-2-1, anterior fragment of an
46, article 40.1), and (2) because the name Indrico- eroded left magnum (Fig.2A–C), housed in the
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592
GIANT RHINOCEROTOID FROM ANATOLIA 583
Figure2. Protaceratherium sp. cf. Protaceratherium albigense (Roman, 1912). Left fragmentary magnum (GK-2-1).
Gözükizilli-2localtity(Çankiri-ÇorumBasin,CentralAnatolia,Turkey).LateOligocene.A,anteriorview;B,medialview;
C,distalview.Scalebar=2cm.Paraceratheriumsp.Rightradius,proximalend(GK-2-2).Gözükizilli-2locality(Çankiri-
Çorum Basin, Central Anatolia, Turkey). Late Oligocene. D, anterior view; E, posterior view; F, antero-proximal view.
Scale bar=10cm.
Natural History Museum of the Maden Tektik ve Description and comparison
Arama (MTA) in Ankara. Gözükizilli-2 loc., Lower The bone is small and slender (anterior transverse
Mbr of the Kizilirmak Fm. (north-central Anatolia). diameter=22.7mm; anterior height=22mm;
Late Oligocene (Ünay etal., 2003; this work). For maximal height c.37mm; McIII-facet transverse
location, see Figure1. diameter=22mm). The surface is quite severely
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592
584 P.-O. ANTOINE ETAL.
Figure3. A, Paraceratherium sp. Right radius, proximal end (GK-2-2) in anterior view. Locus 2, Gözükizilli (Çankiri-
Çorum Basin, Central Anatolia, Turkey); B, Paraceratherium transouralicum. Right radius, detail of the proximal end
(AMNH 26166) in anterior view, after Granger & Gregory (1936: 43, fig.24). Scale bar=10cm.
eroded in places. The bone is broken in front of the fig.48.7). No direct comparison can be made with
posterior tuberosity. In anterior view, the anterior P.betpakdalense (Borissiak, 1938) from the Late
side is square, with a regularly convex distal border, Oligocene to earliest Miocene of the ‘Aral suite’ in
while the proximal, lateral and medial borders are Kazakhstan(deBonisetal.,1997),forwhichnomag-
straight.TheinsertionoftheM.extensordigitorumis num is available.
a thick and wide tubercle which occupies the central
part of the anterior side. It is very prominent in its
FAMILY HYRACODONTIDAE COPE, 1879
latero-distal tip. The dorsal process is slender. When
SUBFAMILY INDRICOTHERIINAE BORISSIAK, 1923
comparedwiththeproximo-distalaxis,itsobliquityis
GENUS PARACERATHERIUM FORSTER-COOPER, 1911
c.40°. In medial view, the dorsal process (for the
PARACERATHERIUM SP.
semilunate) is semicircular (diameter=25mm) and
FIGURES2D–F AND 3
posteriorly displaced. It is separated from the ante-
rior side by a smooth inflection. Due to erosion, the Material available: GK-2-2, proximal fragment of a
medialfacets(forthetrapezoidandthesecondmetac- rightradiusfromalargeadultindividual(Fig.2D–F),
arpal) are not preserved. The distal facet is regularly housedintheNaturalHistoryMuseumoftheMaden
concave, with a large diameter. As an inference, the Tektik veArama (MTA) inAnkara. Gözükizilli-2 loc.,
third metacarpal is likely to bear a wide antero- Lower Mbr of the Kizilirmak Fm. (north-centralAna-
posterior diameter. In lateral view, the articular tolia). Late Oligocene (Ünay etal., 2003; this work).
facets are eroded. In distal view, the facet for the For location, see Figure1.
third metacarpal shows a convex anterior border and
straight subparallel lateral borders. The size, shape Description and comparison
and morphological features of GK-2-1 closely match GK-2-2 is a fragmentary right radius from an adult
those of the magnums referred to Protaceratherium individual (proximal epiphysis and diaphysis fused);
albigense (Roman, 1912) from the early Late Oli- only the proximal third is preserved, including the
gocene of Marseilles, France (P. O. Antoine, pers. proximalpartofthediaphysis.Moredistalfragments
observ.), and, to a lesser degree, with that of P.minu- of the same bone were found in situ, scattered as a
tum (Cuvier, 1822) from the Early Miocene of result of weathering. The bone was lying on its ante-
Laugnac, France (de Bonis, 1973). The latter differs rior side, at the bottom of a conglomerate lens 30cm
byhavinganarroweranteriorsideandtheabsenceof thick. The proximal articulation has furrows, some
aposteriorlydisplaceddorsalprocess(deBonis,1973: over several centimetres in diameter; they may origi-
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GIANT RHINOCEROTOID FROM ANATOLIA 585
Table1. Comparison of measurements of radii (mm) from large indricothere rhinocerotoids, after Pavlova (1922),
Forster-Cooper(1924),TeilharddeChardin(1926),Granger&Gregory(1936),Gromova(1959),andpersonalobservation
by P.-O.A. (Paraceratherium bugtiense: Bugti Hills, Balochistan)
Paraceratherium Paraceratherium ‘Indricotherium’
Paraceratherium sp. bugtiense prohorovi transouralicum
Measurement Gözükizilli (this work) (our unpubl. data) (Borissiak, 1939) (Pavlova, 1922)
Length pres. 490; est. 1200 – 665; 731; 985 960–1220
prox. end
TD pres. 270; est. 310 170 – (>235) 180–342 241-est. 290
APD pres. 210 100 – (>158) 114–205 178-est. 250
Prox. art.
TD pres. 270; est. 280 170–220 ? est. 260
med. hum. fac.
TD pres. 135; est. 160 – 100–193 123
APD >108 105 – (>120) 85–100 121
lat. hum. fac.
TD 135 – 70–146 108
APD pres. 114; est. 140 75 – (>93) 70–84 ?
lat. ulna fac.
H pres. 60; est. 80 – ? ?
min. dia.
TD est. 185 90–125 110–185 147-est. 160
APD pres. 80; est. >100 75–90 70–112 92-est. 110
Baluchitherium grangeri is here considered a junior synonym of Indricotherium transouralicum, as stated by Lucas &
Sobus(1989).ThemeasurementsprovidedbyGranger&Gregory(1936)for‘B.grangeri’areincludedinthecorresponding
column.
APD, antero-posterior diameter; art., articulation; dia., diaphysis; est., estimated; H, height; hum., humerus; min.,
minimal; pres., preserved; prox., proximal; TD, transverse diameter.
nate from post-mortem predation. The proximal phology is characteristic of the Rhinocerotoidea
extremity is not deformed, while the diaphysis dis- among Perissodactyla. Contiguous with the posterior
plays longitudinal cracks and is crushed in its poste- border of the lateral cochlea, a small part of the
rior side. The preserved part is very large (Table1), lateral ulna facet is visible. It is oblique both trans-
with an estimated bone length of c.1200mm (esti- versely and sagittally.
mated by comparison with available radii from other The anterior side is badly damaged in its proximal
indricotheres; Table1). The cortical part (substantia part. In anterior view, the median ridge between the
corticalis) of the diaphysis exceeds 32mm in thick- biconcavehumerusfacetsisverysmooth,anditforms
ness. GK-2-2 displays characteristic proportions of a a dihedral angle of about 135° (Fig.2E). As a conse-
cursorial rhinocerotoid. The proximal end is wide quence, the preserved part of the proximal border of
and deep (transverse diameter c.310mm; antero- the bone is ‘W’ shaped. The anterior side displays a
posterior diameter greater than 210mm), nearly wide and salient axial tubercle, with a rough surface.
three times larger than the mean values observed in This tubercle is located very proximally. It forms an
living black rhinos (Guérin, 1980: 85, table9). It is equilateral triangle (140mm wide) tapering distally
widened laterally with respect to the diaphysis. and corresponding to the insertion for the M.biceps
Inproximalview,thehumerusarticulationshowsa brachii. In most rhinocerotoids, this insertion is flat
pear-like outline, with a concave anterior border and or depressed (Antoine, 2002). Indeed, only indrico-
a larger medial part. It consists of two cochleae occu- there hyracodontids bear such a salient insertion
pying most of the proximal surface. The medial among Rhinocerotoidea (e.g. Granger & Gregory,
cochlea has largely been destroyed while the lateral 1936: fig.24). There is a shallow vertical groove on
one is better preserved (Fig.2F). Both cochleae are the lateral half of the anterior side, externally from
biconcave. The lateral cochlea is much smaller but the insertion of the M.biceps brachii. More distally,
moreconcave.Itshowsatrapezoidaloutlineinproxi- the anterior surface of the diaphysis is smooth and
mal view (Fig.2F) and lacks any lateral extension regular. The proximal articulation and the medial
corresponding to the capitulum humeri. Such mor- border of the diaphysis form an angle of c.100°
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592
586 P.-O. ANTOINE ETAL.
(the medial border is broken proximally). The lateral rium Borissiak, 1915, Aralotherium Borissiak, 1939
border is strongly convex in its proximal part: the andDzungariotheriumChiu,1973.Bycontrast,other
maximal width of the bone (c.310mm) is situated authors(e.g.Gromova,1959;Spassov,1989;Fortelius
c.80mm below the proximal end. In this area is the & Kappelman, 1993) consider Indricotherium and
insertion for the M.extensor digitorum communis Paraceratherium as distinct genera.
(Guérin, 1980: 83–84, fig.11). Distally to this Pending a forthcoming systematic revision of the
maximal width, the diaphysis tapers quickly. This phylogeny of indricotheres, the views of Lucas &
tapering is stronger laterally than medially, the Sobus (1989) are followed: a single Oligocene genus,
lateral border being very oblique from the vertical Paraceratherium, with four valid species – P.bug-
(Fig.2D). tiense(Pilgrim,1910)fromtheBugtiHillsofPakistan
In lateral view, the proximal articulation is much (Pilgrim, 1910; Forster-Cooper, 1913a, b, 1924, 1934;
higherposteriorlythananteriorly,whichisextremely Welcomme etal., 1999, 2001; Antoine etal., 2004);
prominent along the median ridge between the P.transouralicum (Pavlova, 1922) and P.prohorovi
cochleae. This obliquity is similarly observed in most (Borissiak,1939)fromAsiaandsouth-easternEurope
fossil and living rhinos (Guérin, 1980). (Osborn, 1923; Granger & Gregory, 1936; Gromova,
The posterior side of the bone probably suffered 1959; Nikolov & Heissig, 1985; Codrea, 2000); and
from a long period of exposure to erosion and/or P.orgosensis(Chiu,1973)fromChina(Chiu,1973;Xu
post-mortempredation:itisstillmorebadlydamaged & Wang, 1978).
than the anterior side. In posterior view, the highest No direct comparison can be made with Paracer-
part of the bone corresponds to the limit between the atherium orgosensis or Urtinotherium Chow & Chiu,
cochleaeforthehumerus.Themedialulnafacetisnot 1963, nor with the controversial indricothere Benar-
preserved. The lateral ulna facet is wide, as is con- atherium callistrati Gabunia, 1955 from the Late
stantly observed among Rhinocerotoidea. It is pre- Oligocene of Georgia (Gabunia, 1964), because no
served in two places (Fig.2D): (1) a flat and crescent- radius is available for these latter taxa.
like surface near the axis of the bone and contiguous By contrast, GK-2-2 shows no substantial morpho-
to the proximal humerus facet – it is oblique trans- logical difference with the radii of P.bugtiense
verselyandvertically(about50°fromthevertical);(2) recently found in the Late Oligocene of the Bugti
a centimetre-sized surface, 60mm more distally than Hills, Pakistan, and particularly with those from
theformer–itissimultaneouslyobliqueandconcave the type locality of this species (Lundo-Chur J2;
transversally, and flat antero-posteriorly. Distally to Welcomme etal., 2001; Antoine etal., 2004; our
the articulate region, the axial part of the posterior unpubl. data): the surface of the bone is rough, with
side forms a salient ridge which separates two deep averythickparscorticalis;therearetwocochleaefor
and wide depressions. These depressions are covered thehumerus,slightlyasymmetricandseparatedbya
byaroughsurface,indicatingatightcontactwiththe smooth ridge, the lateral one being biconcave; the
ulna by means of a strong membrana interossea. posterior part of the humeral articulation is much
In distal view, the diaphysis is ovoid in cross- higher than the anterior border; the insertion for the
section. M.biceps brachii is wide, salient and triangular; the
Although fragmentary, the specimen from diaphysis tapers strongly distally from the proximal
Gözükizilli-2displaysacombinationofsizeandshape end; and there is a shallow vertical groove on the
that fits only with the radii referred to large indrico- lateral half of the anterior side, externally from the
there rhinocerotoids (Table1): for instance, other insertion of the M.biceps brachii. The specimen from
megamammals such as elephantoids bear reduced Gözükizilli-2islargerthanthelargestradiusreferred
and slender radii with respect to their body size. to P.bugtiense (Table1).
Lucas & Sobus (1989) published the last systematic In addition, GK-2-2 is very similar to the largest
revision of indricotheres, with four valid genera rec- specimen attributed to Baluchitherium grangeri
ognized within the subfamily Indricotheriinae Boris- Osborn,1923byGranger&Gregory(1936:43,fig.24);
siak, 1923: the small-sized genera Forstercooperia the latter is a junior synonym of P.transouralicum
Wood, 1939 and Juxia Chow & Chiu, 1964, mostly accordingtoLucas&Sobus(1989:367).Onametrical
recordedfromEocenelocalitiesinAsia,andthelarge- basis, they are equivalent (Table1; Fig.3). In P.pro-
sizedgeneraUrtinotheriumChow&Chiu,1963,from horovi from the Aral Suite of Kazakhstan, the size
the Early Oligocene of Asia, and Paraceratherium distribution of the radii appears bimodal (Table1;
Forster-Cooper, 1911, from the late Early and Late Gromova, 1959) while the morphology is constant. A
Oligocene ofAsia and eastern Europe. In that sense, similarbimodality,probablyreflectingastrongsexual
Paraceratherium includes all species previously dimorphism,isobservedforthe‘frequencydistribution
referredtoThaumastotheriumForster-Cooper,1913a, of mass estimates’ in P.transouralicum (Fortelius &
Baluchitherium Forster-Cooper, 1913b, Indricothe- Kappelman,1993:96).ThesameoccursinP.bugtiense
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GIANT RHINOCEROTOID FROM ANATOLIA 587
fromPakistan(Antoineetal.,2004).Thisphenomenon in Asia and Eastern Europe (Lucas & Sobus, 1989;
has long been debated for indricotheres, especially Protheroetal.,1989;Lucas,Kordikova&Emry,1998;
concerning its taxonomic implications (i.e. Lucas & Meng & McKenna, 1998; Welcomme etal., 2001;
Sobus, 1989 vs. Spassov, 1989; Fortelius & Kappel- Antoineetal.,2004).Unfortunately,nomorphological
man, 1993; Antoine etal., 2004). evolution can be ascertained for the whole interval of
Therefore,andpendingthediscoveryofsupplemen- thegenus–especiallyonpostcranials–whichdoesnot
tary material, we propose to refer the indricothere allowfurtherdirectprecisionregardingtheageofthe
from Gözükizilli-2 to Paraceratherium sp., further Gözükizilli-2locality.Yet,aLateOligoceneagecanbe
presuming that the radius belongs to a large indi- inferred for Gözükizilli-2 thanks to its stratigraphic
vidual, i.e. most probably to a male. position, intercalated between a Late Oligocene local-
ity (Gözükizilli-1 in the Lower Mbr) and two Late to
latest Oligocene loci (Tepe 641 and Kizilirmak in the
AGE OF THE KIZILIRMAK FORMATION Upper Mbr; see below).
Priorto2002,asinglemammallocalitywasknownin
the Kizilirmak Fm. (Gözükizilli-1; Ünay etal., 2003). TEPE 641 (FIG.1C)
Several mammal localities were discovered within
The provisional mammal list is:
the Kizilirmak Fm. during the 2002 fieldwork
(Gözükizilli-2 in the Lower Mbr; Tepe 641 and – Iberomeryx cf. parvus Gabunia, 1964 [majority
Kizilirmak in the Upper Mbr). The corresponding among artiodactyls],
faunas(listedbelow)allowustoinferatimerangefor – Tragulidae indet. sp. 1
this formation. – ?Palaeohypsodontus sp.,
– Cervoidea gen. et sp. indet. (size of Procervulus/
Dremotherium),
GÖZÜKIZILLI-1 (FIG.1C) – ?Feliformia indet. (small),
The faunal list provided by Ünay etal. (2003) is as – Rhinocerotidae gen. et sp. indet.,
follows, with familial assignment in square brackets – Rhinocerotoidea gen. et sp. indet.,
for the taxa identified at the species/genus level: – Rodentia, gen. et sp. indet.,
– Lagomorpha, gen. et sp. indet.
– aff. Sayimys sp. [Ctenodactylidae],
– Ctenodactylidae gen. B. sp. 1,
– Dipodidae indet., KIZILIRMAK (FIG.1C)
– Eucricetodon sp. 2 [Cricetidae], The provisional faunal mammal list is:
– Eucricetodon sp. 3 [Cricetidae],
– Iberomeryx cf. parvus Gabunia, 1964,
– Glirulus sp. [Gliridae],
– Tragulidae indet. sp. 1 [same as that of Tepe 641;
– Bransatoglis cf. complicatus [Gliridae]
majority among artiodactyls]
A similar assemblage, with ctenodactylids and crice- – Carnivora indet., sp. 1,
tids, is reported in the Central Anatolian locality of – Carnivora indet., sp. 2,
Yeniköy, assumed to be coeval with the Gözükizilli – Rhinocerotidae gen. et sp. indet.,
site (Ünay etal., 2003). – Rodentia, gen. et sp. indet.,
– Lagomorpha, gen. et sp. indet.
GÖZÜKIZILLI-2 (FIG.1C) A coeval age can be hypothesized for Tepe 641 and
Althoughpoorlydiversifiedandimpreciselyidentified, Kizilirmak,basedonbothlithostratigraphy(floodplain
both rhinocerotoids provide a rather precise age for depositsintheUpperMbroftheKizilirmakFm.)and
Gözükizilli-2:therhinocerotidProtaceratheriumhasa biostratigraphy. The difference between these locali-
goodrecordfromthelateEarlyOligocenetotheEarly ties (e.g. in the relative abundance of artiodactyls) is
Miocene in Eurasia (MP24 to MN 4; Marivaux, 1999; probablyduetotaphonomicalbias(mostremainswere
Antoine,Bulot&Ginsburg,2000;Antoineetal.,2003). damaged by predators or scavengers).
The species showing the closest affinities with the In Tepe 641, the small and slender Iberomeryx cf.
currently described specimen is P.albigense (Roman, parvus is represented by 20 cheek teeth (and dozens
1912), which has an MP 24–28 stratigraphic range of isolated postcranials) with typical lophiomerycid
(Uhlig, 1999; Antoine etal., 2003). Moreover, this features, such as the lingual opening of the anterior
species has recently been described in late Early to and posterior fossettes on lower molars (Janis, 1987:
early Late Oligocene coastal deposits from Turkish 214, fig.10). Both the lack of any parastyle on the
Thrace (Saraç, 2003). The giant rhino Paracerath- upper molars and the backward elongation of
eriumoccursfromthelateEarlytothelatestOligocene the metaconid on the fourth premolar distinguish
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592
588 P.-O. ANTOINE ETAL.
the genus Iberomeryx, as demonstrated by Métais (2003: 98) states that, during the Oligocene, ‘in Asia
etal. (2001). This genus is known throughout the [the strong endemic development of rodents] can be
Oligocene in Eurasia (Gabunia, 1964; Sudre, 1984; seen most clearly in the array of ctenodactyloids and
Janis, 1987; Nanda & Sahni, 1990; Sudre & Blondel, in Europe of glirids and theridomyids’. In fact, the
1996; Métais etal., 2001). The closest affinities exist Oligocene spatial distribution of Ctenodactylidae is
with the type species, I.parvus, from the Late Oli- limited westward to Anatolia (de Bruijn, 1999;
goceneofGeorgia(Gabunia,1964,1966;Sudre,1984). Dawson, 2003). Among this poor Anatolian record,
However,IberomeryxfromTepe641seemstobemore Gözükizilli is the westernmost locality where this
advanced than any known species of the genus, espe- endemic Asian rodent group occurs (Ünay etal.,
cially in possessing very elongate lower molars and a 2003). At the same time, Gözükizilli-1 is the first
strong lingual cingulum on all the upper molars. Oligocene glirid-bearing locality (Ünay etal., 2003),
The recognition of ?Palaeohypsodontus sp. is based as the Gliridae were previously restricted to
on two fragmentary molars. These teeth are as hyp- European Oligocene localities. Similarly, some other
sodont as bovid teeth, but their structures are very Anatolian Oligocene localities have yielded the
simple.Astate-of-the-art revision of Palaeohypsodon- anthracotheres Anthracotherium and Elomeryx,
tus has recently been made (Métais etal., 2003), which have a wide Eurasian distribution (Russell
showing that this puzzling genus was a hypsodont etal., 1982; Saraç, 2003).
traguloid rather than a precocious bovid. Indeed, The Late Oligocene mammal faunas from Anatolia
unambiguous remains of Palaeohypsodontus are share several elements with more or less coeval
restricted to two localities from Mongolia (Early and faunas from Pakistan and Mongolia (Paracerath-
Late Oligocene; Vislobokova & Daxner-Höck, 2002) erium, Palaeohypsodontus and ctenodactylids;
and the Lundo-Chur J2 locality in Baluchistan Dashzeveg, 1990; Métais etal., 2003; Antoine etal.,
(Late Oligocene; Métais etal., 2003). The occurrence 2004), Georgia (Iberomeryx and an indricothere),
of Palaeohypsodontus in Central Anatolia would Central Europe (Anthracotherium and Paracerath-
widely expand westward the geographical range of erium) and Western Europe (Protaceratherium,
this puzzling ruminant genus. Anthracotherium and glirids). In summary, these
The morphological advancement of Iberomeryx strong affinities with bothAsia (including the Indian
tends to indicate a latest Oligocene age for Tepe 641, subcontinent) and Europe do not support any island
as Iberomeryx and Palaeohypsodontus have a strictly hypothesis for the Late Oligocene of Anatolia. In
Oligocene record (Métais etal., 2003). contrast,themammalfaunasfromtheLateOligocene
Kizilirmak Fm. of north central Anatolia have not
thus far yielded any taxon of Arabo-African affinity.
PALAEOGEOGRAPHY
On an environmental point of view, ctenodactyloid
Among the mammals recognized in the Kizilirmak rodents (Gözükizilli-1; Lower Mbr) and hypsodont
Fm., three groups (rhinocerotoids, ruminants, herbivores such as ?Palaeohypsodontus and lagomor-
rodents) allow us to enhance the palaeogeographical phs (Tepe 641 and Kizilirmak; Upper Mbr) occur
relationships of north Central Anatolia during the mainly in Shandgolian (Early Oligocene) and Taben-
Late Oligocene. bulukian (Late Oligocene) localities of Mongolia and
The small rhinocerotid from Gözükizilli-2 has China (McKenna & Bell, 1997; Wang, 1997; Meng &
closest affinities with Protaceratherium albigense McKenna, 1998; Métais etal., 2003). This kind of
fromwesternandcentralEuropeandTurkishThrace assemblage strongly suggests the existence of abra-
(Antoine etal., 2003; Saraç, 2003). Indricotheres are sive vegetation in the surroundings, likely to occur
essentiallyrecordedinAsia(Antoineetal.,2004).Yet, in rather open habitats (Wang, 1997). By contrast,
a few Oligocene localities have yielded scarce indri- the glirids Glirulus and Bransatoglis (Gözükizilli-1;
cothere remains around the Black Sea, in the Cauca- Lower Mbr) were most probably scansorial rodents
sus and Eastern Europe (Fig.4). Indeed, Gözükizilli typical of forest habitats (Hartenberger, 1994), while
is the first indricothere-bearing locality south of the the unidentified tragulid (Tepe 641 and Kizilirmak;
Paratethys(anareamoreorlesscorrespondingtothe Upper Mbr) rather tends to indicate a water-related
recent Black Sea; Fig.4). Among the ruminants rec- forestedenvironment(Rössner,1997;Gentry,Rössner
ognized in the Kizilirmak Fm. deposits, Iberomeryx & Heizmann, 1999).
has a strict westernAsian record (Janis, 1987), while
the occurrence of a probable Palaeohypsodontus
ACKNOWLEDGEMENTS
would widely expand westward the geographical
range of this puzzling ruminant genus, thus fae only The 2002 fieldwork in the Çankiri-Çorum basin was
recognized in Mongolia and Pakistan (Vislobokova & fundedbyTÜBITAK,theCNRS(ECLIPSEProgram),
Daxner-Höck, 2002; Métais etal., 2003). Dawson and the Faculty of Sciences of the University of
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592
GIANT RHINOCEROTOID FROM ANATOLIA 589
Figure4. Palaeogeographical map of Middle East and south-west Asia illustrating the geographical distribution of
Oligocene indricotheres around the Paratethys, in Kazakhstan and in Pakistan. After Petronijevic & Thenius (1957),
Gromova (1959), Gabunia & Iliescu (1960), Gabunia (1964), Nikolov & Heissig (1985), Lucas & Sobus (1989), Spassov
(1989),Lucas&Bayshashov(1996),Lucas&Emry(1996),Lucasetal.(1998),Codrea(2000),Welcommeetal.(2001)and
Antoine etal. (2004). Source for map construction: http://www.odsn.de/odsn/services/palaeomap/adv_map.html
Ankara. Gürol Seyitoglu and Baki Varol (Fen Fakül- Antoine PO, Ducrocq S, Marivaux L, Chaimanee Y,
tesi, Jeoloji Mühendisligi Bölümü,Ankara),Arzu Gül Crochet JY, Jaeger JJ, Welcomme JL. 2003. Early
(Denizli University), Raquel Lopez Antonanzas and rhinocerotids (Mammalia, Perissodactyla) from South Asia
FabienKnoll(MuséumNationald’HistoireNaturelle, andareviewoftheHolarcticPaleogenerhinocerotidrecord.
Paris) contributed to this field trip. Grégoire Métais Canadian Journal of Earth Sciences 40: 365–374.
providedusefulliteraturesources(andadvice)forthe Antoine PO, Shah SMI, Cheema IU, Crochet JY, de
identification of the artiodactyls from Tepe 641 and Franceschi D, Marivaux L, Métais G, Welcomme JL.
2004. New remains of the baluchithere Paraceratherium
Kizilirmak.Thisarticleisdedicatedtothememoryof
bugtiense (Pilgrim, 1910) from the Late/latest Oligocene of
Annie Antoine.
the Bugti Hills, Balochistan, Pakistan. Journal of Asian
Earth Sciences 24: 71–77.
de Bonis L. 1973. Contribution à l’étude des mammifères
REFERENCES
de l’Aquitanien de l’Agenais. Rongeurs-Carnivores-
AbelO.1910.KritischeUntersuchungenüberdiepaläogenen Périssodactyles. Mémoires du Muséum National d’Histoire
Rhinocerotiden Europas. Abhandlungen der Geologische Naturelle, Paris 28: 1–192.
Reichsanstall, Wien 20: 1–52. deBonisL,BrunetM,KordikovaEG,MavrinAV.1997.
Antoine PO. 2002. Phylogénie et évolution des Elasmoth- Oligocene–Miocene sequence stratigraphy and vertebrate
eriina (Mammalia, Rhinocerotidae). Mémoires du Muséum paleontology of western and southern Betpakdala Steppe,
National d’Histoire Naturelle, Paris 188: 1–359. SouthKazakhstan.In:AguilarJP,LegendreS,MichauxJ,
AntoinePO,BulotC,GinsburgL.2000.Unefaunerarede eds.ActesduCongrèsBiochroM’97.MémoiresetTravauxde
rhinocérotidés(Mammalia,Perissodactyla)dansleMiocène l’Ecole pratique des Hautes Etudes, Institut de Montpellier
inférieurdePellecahus(Gers,France).Geobios33:249–255. 21: 225–240.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592
590 P.-O. ANTOINE ETAL.
Borissiak AA. 1915. Ob indrikoterii (Indricotherium n.g.). Forster-Cooper C. 1934. The extinct Rhinoceroses of
Geologiskie Vestnik 1: 131–134. Baluchistan.PhilosophicalTransactionsoftheRoyalSociety
Borissiak AA. 1923. Sur un nouveau représentant des rhi- of London, Series B 223: 569–616.
nocéros gigantesques de l’Oligocène d’Asie. Indricotherium ForteliusM,KappelmanJ.1993.Thelargestlandmammal
asiaticum, n.g. n.sp. Mémoires de la Société Géologique de ever imagined. Zoological Journal of the Linnean Society
France 59: 1–16. 108: 85–101.
Borissiak AA. 1938. K filogenii Dicerorhinae. Doklady Aka- Gabunia L. 1955. O svoebraznom pred stavitele Indri-
demii Nauk SSSR 19: 767–770. cotheriidae iz Oligotsena Gruzii. Doklady Akademie Nauk
BorissiakAA.1939.Onovompredtavitelecem.Indricotheri- Armeniya SSSR 21: 177–181.
idae. In: V.A. Obruchevu k 50letiyu nauchoj i pedagog- Gabunia L. 1964. Benarskaya Fauna Oligotsenovykh poz-
icheskoj deyatelnosti. Isdatelskovo Akademia Nauk SSSR, vonochnykh. Tbilisi: Metsniereba Press. [in Russian, with
Moscow 2: 271–276. Georgian and French abstracts]
de Bruijn H. 1999. Superfamily Ctenodactyloidea. In: GabuniaL.1966.SurlesMammifèresoligocènesduCaucase.
RössnerGE,HeissigK,eds.TheMiocenelandmammalsof BulletindelaSociétégéologiquedeFrance7:857–869.
Europe. Munich: Pfeil Verlag, 263–266. Gabunia L, Iliescu O. 1960. O pervoij nakhodke ostatkov
ChiuC.1973.Anewgenusrhinoceros[sic]fromOligoceneof gigantiskikh nosorogoviz sem. Indricotheriidae v Rumynii.
Dzungaria, Sinking. Vertebrata PalAsiatica 11: 182–191. Doklady Akademia Nauk SSSR 130: 25–427.
ChowM,ChiuC.1963.Anewgenusofgiantrhinocerosfrom Gentry AW, Rössner GE, Heizmann EPJ. 1999. Suborder
OligoceneofInnerMongolia.VertebrataPalAsiatica7:230– Ruminantia. In: Rössner GE, Heissig K, eds. The Miocene
239. land mammals of Europe. Munich: Pfeil Verlag, 225–258.
Chow M, Chiu C. 1964. An Eocene giant rhinoceros. Verte- GrangerW,GregoryWK.1936.Furthernotesonthegigan-
brata PalAsiatica 8: 264–267. tic extinct rhinoceros, Baluchitherium, from the Oligocene
Codrea V. 2000. Rinoceri si Tapiri Tertiari din România. of Mongolia. Bulletin of the American Museum of Natural
Cluj: Presa Universitara Clujeana. History 72: 1–73.
Cope ED. 1879. On the extinct Species of Rhinoceridae of Gray JE. 1821. On the natural arrangements of vertebrose
NorthAmericaandtheirallies.BulletinoftheUnitedStates animals. London Medical Repository 15: 296–310.
Geological and Geographic Survey 5: 1–227. Gromova V. 1959. Gigantskie nosorogi. Trudy Paleontology
CuvierG.1822.Recherchessurlesossemensfossiles,oùl’on Institut Akademii Nauk SSSR 71: 1–164.
rétablitlescaractèresdeplusieursanimauxdontlesrévolu- GuérinC.1980.LesRhinocéros(Mammalia,Perissodactyla)
tionsduglobeontdétruitlesespèces.4èmeédition.Paris,10 du Miocène terminal au Pléistocène supérieur en Europe
vol. occidentale. Comparaison avec les espèces actuelles. Docu-
Dashzeveg D. 1990. The earliest rodents (Rodentia, Cteno- ments du Laboratoire de Géologie de l’Université de Lyon,
dactylidae) of Central Asia. Acta Zoologica Cracoviensis, Sciences de la Terre 79: 1–1184.
Krakow 33: 11–35. Hartenberger JL. 1994. The evolution of the Gliroidea. In:
DawsonMR.2003.PaleogenerodentsofEurasia.In:Reumer TomidaY,LiCK,SetoguchiT,eds.RodentandLagomorphs
JWF,WesselsW,eds.DistributionandmigrationofTertiary of Asian origin and diversification. National Science
mammals in Eurasia. Rotterdam: DEINSEASpecial Issue, Museum Monographs, Tokyo 8: 19–33.
97–126. Janis CM. 1987. Grades and clades in hornless rumniant
Forster-Cooper C. 1911. Paraceratherium bugtiense, a new evolution: the reality of the Gelocidae and the systematic
genusofRhinocerotidaefromtheBugtiHillsofBaluchistan. position of Lophiomeryx and Bachitherium. Journal of Ver-
Preliminarynotice.AnnualMagazineofNaturalHistory8: tebrate Paleontology 7: 200–216.
711–716. KaradenizliL,SaraçG,SenS,AntoinePO,SeyitogluG,
Forster-CooperC.1913a.Thaumastotheriumosborni,anew Varol B, Hakyemez Y. 2003. Türkiye’de Bir Iik: Bütün
genusofperissodactylesfromtheUpperOligocenedeposits Zamanlarin En Büyük Karasal Memeli Hayvani Baluchi-
of the Bugti Hills of Baluchistan. Preliminary notice. therium (Paraceratherium) Çankiri-Çorum Havzasinda
Annual Magazine of Natural History 12: 376–381. Bulundu [A first in Turkey: Baluchitherium (Paracerath-
Forster-Cooper C. 1913b. Correction of generic name erium),theeverlargestlandmammalofalltimes,hasbeen
[Thaumastotherium to Baluchitherium]. Annual Magazine found in Çankiri-Çorum Basin], 56th Turkish Geological
of Natural History 12: 504. Congress,Ankara, 14–20thApril 2003, MTA, 292–294.
Forster-Cooper C. 1915. New genera and species of Lucas SG, Bayshashov BU. 1996. The giant rhinoceros
mammals from the Miocene deposits of Baluchistan. Pre- Paraceratherium from the late Oligocene at Aktau Moun-
liminary notice. Annual Magazine of Natural History 16: tain,southeasternKazakhstan,anditsbiochronologicalsig-
404–410. nificance. Neues Jahrbuch für Geologie und Paläontologie,
Forster-Cooper C. 1924. On the skull and dentition of Mitteilhungen, Stuttgart 9: 539–548.
Paraceratherium bugtiense: a genus of aberrant rhinoceros Lucas SG, Emry RJ. 1996. Early record of indricothere
fromthelowerMiocenedepositsofDeraBugti.Philosophi- (Mammalia: Perissodactyla: Hyracodontidae) from theAral
cal Transactions of the Royal Society of London, Series B Sea region of western Kazakhstan. Proceedings of the Bio-
212: 369–394. logical Society of Washington 109: 391–396.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 581–592