Late Miocene bovids from Şerefköy-2 (SW Turkey)
and their position within the sub-Paratethyan
biogeographic province
DIMITRIS S. KOSTOPOULOS and SEVAL KARAKÜTÜK
Kostopoulos, D.S. and Karakütük, S. 2015. Late Miocene bovids from Şerefköy-2 (SW Turkey) and their position within
the sub-Paratethyan biogeographic province. Acta Palaeontologica Polonica 60 (1): 49–66.
We describe new fossil bovid craniodental remains from the Upper Miocene fossil site of Şerefköy-2, Yatağan Basin,
SW Turkey. The new material belongs to six species: Gazella cf. G. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, Urmiatherium rugosifrons, and ?Sinotragus sp., which together indicate a latest
middle–early Late Turolian (Late Miocene) age. Medium-to-large bovid taxa prevail over small ones, and protoryxoid
bovids clearly dominate the assemblage. An analysis of the taxonomic structure, size and diet spectra of several Turolian bovid assemblages from Greece and Turkey reveals Şerefköy-2 to be a member of a mammalian palaeocommunity
particular to southwestern Anatolia, which in turn forms part of the sub-Paratethyan biogeographic province.
Ke y w o rd s : Mammalia, Bovidae, Antilopinae, biogeography, Miocene, Turolian, Turkey, Anatolia.
Dimitris S. Kostopoulos [dkostop@geo.auth.gr], Aristotle University of Thessaloniki, School of Geology, 54124 Thessaloniki, Greece.
Seval Karakütük [sevaloruc.ege.univ@hotmail.com], Natural History Museum, Ege University, 35100 Izmir, Turkey.
Received 7 November 2012, accepted 10 June 2013, available online 17 June 2013.
Copyright © 2015 D.S. Kostopoulos and S. Karakütük. This is an open-access article distributed under the terms of the
Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium,
provided the original author and source are credited.
Introduction
Anatolia, located in modern-day Turkey, plays a crucial role in
the reconstruction of late Neogene Eurasian palaeozoogeography and palaeoenvironments (e.g., Costeur and Legendre 2008;
Eronen et al. 2009; Kostopoulos 2009b), because it represents
the natural link between the western and eastern extremes of the
so-called Late Miocene sub-Paratethyan biogeographic province (Bernor et al. 1979). There area is rich in Late Miocene
fossil sites, but in most cases their contents have only been
reported in the form of preliminary faunal lists (e.g., Sickenberg et al. 1975; Alan 1997; Saraç 2003). Pioneer works by
Ozansoy (1965), Tekkaya (1973a, b), Senyürek (1952, 1953),
Boscha-Erdbrink (1978), and Köhler (1987) only gave a fragmentary view of the rich and abundant Late Miocene bovid assemblage of Anatolia, which were partially, but never entirely,
filled by later studies on newly discovered or re-examined material (e.g., Bouvrain 1994; Gentry 2003; Kostopoulos 2005;
Geraads and Güleç 1999; Bibi and Güleç 2008).
Thee field seasons at the new locality of Şerefköy-2, discovered in the summer of 2007 in Yatağan Basin, SW Turkey
(Fig. 1; coordinates: N 37°21’47.80’’, E 28°14’11.10’’; see
Kaya et al. 2012 for further information on the geological
context), yielded more than 1200 identifiable fossil speciActa Palaeontol. Pol. 60 (1): 49–66, 2015
mens belonging to more than 25 species of mammals. Together, these represent one of the richest Turolian faunas from
Anatolia, and offer a chance to (i) test faunal correlations
within the sub-Parathethyan biogeographic province and (ii)
further develop the local biochronological framework. Here,
we provide a detailed description of the bovids forming part
of the Şerefköy-2 bovid assemblage, preliminary reported by
Kaya et al. (2012). The remaining taxa recovered from this
site will be described elsewhere.
Institutional abbreviations.—AMNH, American Museum
of Natural History, New York, USA; MYŞE PV, collection
of the Natural History Museum of Ege University, Izmir,
Turkey; KNUA, Kapodistrian National University of Athens, Greece; LGPUT, Laboratory of Geology and Palaeontology, Aristotle University of Thessaloniki, Thessaloniki,
Greece; MNHN, Museum national d’Histoire naturelle, Paris, France; NHMA, Aegean Museum of the Natural History-Zimalis Foundation, Samos Island, Greece; NHMW;
Naturhistorisches Museum Wien, Austria.
Other abbreviations.—APD, anteroposterior diameter; CFA,
correspondence analysis; L, length; n, number of specimens;
TD, transverse diameter; W, width. Upper and lower case
letters denote upper and lower teeth, respectively.
http://dx.doi.org/10.4202/app.2012.0129
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
A
B
Black Sea
Milet
Fm.
50
Ankara
TURKEY
Mediterranean Sea
marl
breccia
Eskihisar Fm.
silty marl
conglomerates
ˆ
fossil site Şerefköy-2
coal deposits
Yatagan Fm.
Muğla
C
This allowed us to (i) overcome differing taxonomic opinions;
(ii) incorporate genera with low abundances; and (iii) emphasise general patterns in taxonomic structure.
Finally, we compared the results the CFA analyses with
the size distribution and diet spectra of the involved bovid
faunas. The latter reflect crucial ecological parameters (body
size and feeding prefeences) that are relevant at the level
of both the organism and the community as a whole (e.g.,
Western 1979; McNaughton and Georgiadis 1986; Eisenberg
1990). Multivariate statistics were performed using PAST
version 2.16 (Hammer et al. 2001). Skull and horn core nomenclature follows Gentry (1971, 1992). Dental nomenclature follows Bärmann and Rössner (2011). All measurements
are in millimetres (mm). Further details on all the analyses
and the data themselves are provided as Supplementary Online Material (SOM 1, SOM 2, available at http://app.pan.pl/
SOM/app60-Kostopoulos_Karakutuk_SOM.pdf).
Systematic palaeontology
Class Mammalia Linnaeus, 1758
Order Artiodactyla Owen, 1848
Family Bovidae Gray, 1821
Subfamily Antilopinae Gray, 1821 (including all nonBovinae taxa sensu Kingdon 1982; e.g., Groves and Grubb 2011)
Genus Gazella Blainville, 1816
Type species: Gazella dorcas Linnaeus, 1758, Recent.
Gazella cf. G. capricornis (Wagner, 1848)
Fig. 2.
D
Fig. 1. The fossil mammal site of Şerefköy-2 (Turkey). A. Geographical location. B. Local lithostratigraphy (modified from Kaya et al. 2012). C. Typical fossil accumulation at Şerefköy-2. D. View of the fossiliferous site.
Material and methods
Horn core and cranial proportions of different bovids were explored using discriminant and Principal Components analyses.
Following Bibi and Güleç (2008), Kostopoulos (2009b), Koufos et al. (2006, 2009a), and Kostopoulos and Bernor (2011),
we performed a correspondence analysis (CFA) to investigate
the structure of various Turolian (8.7–5.4 Ma; Late Miocene)
bovid assemblages from Greece, Turkey and Iran. Following
Bibi and Güleç (2008), we first analysed bovid assemblages
according to their relative abundances at the genus level. Next,
we repeated the analysis by grouping all genera into five units
reflective of the basic taxonomic structure of each assemblage.
Material.—MYŞE PV-2501, frontlet; MYŞE PV-1577, PV1834, right horn cores; MYŞE PV-560, PV-1578, left horn
cores; MYŞE PV-1831, PV-1832, partial right horn cores;
MYŞE PV-1359, PV-1632, PV-1828, partial left horn cores;
MYŞE PV-1360, partial horn core; MYŞE PV-1830, 1633,
1393, distal portions of horn cores; MYŞE PV-2572, palate;
MYŞE PV-2575 right upper tooth row P2–M3; MYŞE PV2000, PV-2558, PV-2562, PV-2564, right mandibular body
with p2–m3; MYŞE PV-2557, PV-2563, left mandibular
body with p2–m3; MYŞE PV-2565, left mandibular body
with p3–m3; MYŞE PV-1526, right m3. All from Şerefköy-2,
Turkey, Late Turolian (Late Miocene).
Description.—Gazella cf. G. capricornis is the only gazelle
present at Şerefköy-2, and represented by at least 5 individuals. The supraorbital foramen is located close to the pedicle,
within an elliptical and rather narrow depression (Fig. 2C1).
The postcornual fossa is oval, small, and moderately deep.
The pedicle is moderately high anteriorly (Fig. 2C). The horn
cores are short (maximum length: 100 mm along the anterior
surface), inserted moderately far apart from each other, and
weakly divergent distally (Fig. 2C1). In lateral view, the horn
core is slightly curved and moderately inclined posteriorly
(Fig. 2A, B, C2), while in cross section it changes from be-
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
A
B
51
C1
C2
5 cm
(A–C)
D1
E
5 cm
D2
(D–F)
F
Fig. 2. The bovid artiodactyl Gazella cf. G. capricornis (Wagner, 1848) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Left horn core
(MYŞE PV-1578) in lateral view. B. Left horn core (MYŞE PV-560) in lateral view. C. Frontlet (MYŞE PV-2501) in frontal (C1) and lateral (C2) views. D.
Left mandibular body (MYŞE PV-2557) in occlusal (D1) and lingual (D2) views. E. Right mandibular body (MYŞE PV-2564) in occlusal view. F. Right
P2-M3 of the palate (MYŞE PV-2572) in occlusal view.
ing symmetrically elliptical near the base (owing to weak
mediolateral compression) to a more rounded shape towards
the tip (TD × 100/APD = 70.4–73.2 at the base, n = 4, and
84.6–104.6 at 7 cm above the base, n = 3; Table 1). Strong,
deep, and continuous furrows run along both the anterior and
posterior surfaces of the horn core.
The premolars are moderately long compared to the molars (Fig. 2D–F), with the upper and lower premolar/molar
ratios equalling about 71% (n = 1) and 54–60% (n = 6),
respectively (SOM 3: Tables 1, 2). There is no entostyle on
the upper molars. M3 has a strong metastyle and a weak mesostyle (Fig. 2F). The anterior conid of p3 and p4 is weak, but
separated from the anterior stylid (MYŞE PV-2000). The meTable 1. Horn core measurements (in mm) of Gazella cf. capricornis
from Şerefköy-2 (Turkey). TD, transverse diameter at the base and at
7 cm above the base; APD, anteroposterior diameter at the base and at
7 cm above the base.
Specimen
MYŞE PV-2501
MYŞE PV-560
MYŞE PV-1577
MYŞE PV-1578
TDbase
16.4
17.8
15.3
16.4
APDbase
23.1
24.3
21.7
22.2
TD7
12.8
14.7
10.0
APD7
14.8
14.1
11.8
solingual conid is elongated and oriented posteriorly on p3,
but more triangular (during early wear) and located relatively
further posteriorly on p4 (Fig. 2D, E). The anterior valley of
p3 and p4 is widely open (Fig. 2D, E). The posterior valley of
p4 is open in the single unworn specimen (MYŞE PV-2000)
and still visible even at advanced stages of wear (Fig. 2D, E).
A small ectostylid is present on m1. The mesostylids on m2
and m3 are strong.
Remarks.—The gazelle from Şerefköy-2 is considerably larger than Gazella ancyrensis Tekkaya, 1973 from Middle Sinap,
as well as G. cf. ancyrensis from Maragheh (Iran), Samos
(Greece), and Kemiklitepe-D (Turkey), but smaller than Gazella mytilinii Pilgrim, 1926 from Samos (Bouvrain 1994;
Kostopoulos 2009a; Kostopoulos and Bernor 2011). Gazella pilgrimi Bohlin, 1935 from Samos and the Axios Valley
(Greece) and Akkaşdağı (Turkey) differs from the Şerefköy-2
specimens in having a shorter pedicle, as well as a longer,
more mediolaterally compressed, and more strongly inclined
horn core (Bouvrain 1996; Kostopoulos 2005, 2009a).
The overall shape of the horn core of the Şerefköy-2
gazelle recalls both Gazella deperdita (Gervais, 1847) from
Western Europe and the SE European Gazella capricornis
(Wagner, 1848). However, the horn core of G. deperdita
52
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
Palaeoryx pallasi (Wagner, 1857)
Discriminant function:
D = -0.75TDbase + 0.21APDbase - 1.14TD7 + 1.31APD7
Fig. 4.
18
G.caprinornis, Pikermi: #55
G. deperdita, Luberon: #31
Hotelling t2: 108.53
P (same): 6.1E-14
16
Frequency
14
12
10
8
6
4
2
0
-8
-6.4
-4.8
-3.2
-1.6
0
1.6
3.2
4.8
6.4
Discriminant
Fig. 3. Discriminant analysis of the horn core proportions of Gazella deperdita from Luberon (= Cucuron), Late Miocene, SW Europe and Gazella
capricornis from Pikermi, Late Miocene, Greece, based on four variables:
TD at the base (TDbase), APD at the base (APDbase), TD at 7 cm above
the base (TD7) and APD at 7 cm above the base (APD7). Arrows indicate
values for the Şerefköy-2 gazelle according to the discriminant function.
Abbreviations: APD, anteroposterior diameter; TD, transverse diameter.
is larger (Fig. 3), much more curved in lateral view, more
inclined posteriorly, and more mediolaterally compressed
along its distal portion. Gazella capricornis was originally described from Pikermi (Greece), but similar forms are
widely distributed between the Balkans and Iran (Kostopoulos 2005, 2009a; Kostopoulos and Bernor 2011). Both Gazella cf. G. capricornis from Samos (particularly specimens
from Mytilinii-1A) and the Şerefköy-2 gazelle are characterised by an anterior conid distinct from the anterior stylid
on p3 and p4, and the closing of the posterior valley of p4 at
late wear stages. Together with G. cf. G. capricornis from
Akkaşdağı and Maragheh, they further differ from typical
specimens of G. capricornis from Pikermi in having slightly
shorter premolars compared to the molars (Table 2).
Genus Palaeoryx Gaudry, 1861
Type species: Antilope pallasi (Wagner, 1857), Pikermi, Greece; Late
Miocene.
Material.—MYŞE PV-2573, right upper tooth row with P2–
M3; MYŞE PV-1293, right upper tooth row with P3–M3;
MYŞE PV-1294/B, partial left upper tooth row with M2–M3;
MYŞE PV-1295, partial left upper tooth row with P4–M1;
MYŞE PV-1599, left mandibular body with p2–m3; MYŞE
PV2574, right mandibular body with p2–m3; MYŞE PV2552, partial of right mandibular body with m2–m3. All from
Şerefköy-2, Turkey, Late Turolian (Late Miocene).
Description.—Specimens from Şerefköy-2 referable to this
taxon are limited to some upper and lower tooth rows, representing at least two individuals. The premolars are long
compared to the molars (Fig. 4, SOM 3: Tables 1, 2), with
upper and lower premolar/molar ratios of 70.8% (n = 1) and
64.3–66.7% (n = 2), respectively. P4 is narrower than both P3
and P2, and P2 is shorter than P3. The anterior style is weakly
developed on P2, but sharp, posteriorly curved and joined to
the anterolabial cone on P3 (Fig. 4C). The “metaconule” is
barely recognisable on P2 and P3, which have a distinctly
convex lingual wall, and absent on P4, in which the anterolingual and posterolingual crista of the lingual cone are
symmetrically developed (Fig. 4C). The anterior style of P4
is more pronounced than the anterolabial cone, with the latter
having a more central position on the labial wall of the tooth
than on P3 (Fig. 4C). All upper molars bear a fossetta (central
islet) and low, double entostyles, originating from both the
protocone and the metaconule (Fig. 4C). The anterior lobe
of M1 and M2 is narrower than the posterior one. The development of the mesostyle on the upper molars ranges from
weak in MYŞE PV-2573 to strong in PV-1293 and PV-1294.
The p2 has a simple and strong anterior stylid and no
anterior conid. The mesolingual conid is also simple, short
and placed both anteriorly and almost perpendicularly to
the longitudinal axis of the tooth (Fig. 4A, B). The posterior
stylid on p3 is much less developed than the posterolingual
conid, with both structures becoming fused during late stages of wear (Fig. 4A, B). The anterior conid and the anterior
stylid are indistinct, thus forming a robust anterior cuspid.
The mesolingual conid is elongated and slants distally, joining the talonid at advanced stages of wear (Fig. 4A). The p4
resembles p3, but is larger, and retains and open posterior
valley until the onset of late stages of wear (Fig. 4A, B). The
posterolabial conid is indistinct on p2 and p3, and weakly
developed on p4. The lower molars have a strong mesostylid,
Table 2. Comparison of Gazella cf. G. capricornis from Şerefköy-2 with G. cf. G. capricornis from different localities and with G. capricornis
from Pikermi. Data are from Kostopoulos (2005, 2009a) and Kostopoulos and Bernor (2011). APD, anteroposterior diameter; L M/m, length of
upper/lower molar row; L P/p, length of upper/lower premolar row; TD, transverse diameter at the base and at 7 cm above the base.
APD × 100/TDbase
APD × 100/TD7
LP/LM × 100
Lp/Lm × 100
Gazella cf. G. capricornis
Şerefköy-2 (Turkey) Akkaşdağı (Turkey) Samos (Greece)
135.4–141.8
103.3–137.6
102.6–139.7
95.9–118.0
100.0–128.9
100.0–125.0
71.1–71.2
67.7–74.6
67.0–73.0
54.0–60.2
55.2–56.4
58.0
Maragheh (Iran)
112.2–147.4
103.8–128.7
76.5
58.8–61.8
Gazella capricornis
Pikermi (Greece)
102.5–139.2
85.0–136.4
75.4–78.4
59.7–70.8
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
53
A
B
5 cm
C
Fig. 4. The bovid artiodactyl Palaeoryx pallasi (Wagner, 1848) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Left mandible (MYŞE PV1599) in occlusal view. B. Right mandible (MYŞE PV-2574) in lingual view. C. Right upper tooth row (MYŞE PV-2573) in occlusal view.
moderate ectostylids (basal pillars), and no anterior cingulid
(goat fold). The third lobe of m3 is rather large and bears a
flat entoconulid, as well as a bulbous hypoconulid (Fig. 4A).
Remarks.—Palaeoryx Gaudry, 1861 is mainly known from
Turolian sites of the sub-Paratethyan zoogeographic province
(e.g., Kostopoulos and Bernor 2011 and literature therein),
although it may have appeared during the Vallesian (e.g., Vislobokova 2005). In Turkey, records of the genus are restricted
to the localities of Kayadibi, Eski Bayırköy, and Mahmutgazi
(Köhler 1987), as well as Kemiklitepe-A, B and Akkaşdağı
(Bouvrain 1994; Kostopoulos 2005). Kostopoulos (2005,
2009a) recognised two species in the Aegean region, P. pallasi and P. majori Schlosser, 1904, which differ from each other
mainly in terms of their cranial and horn core morphology. In
addition, a larger species or variety might exist in Maragheh
(Iran) and Ukraine (Kostopoulos and Bernor 2011).
The specimens from Şerefköy-2 differ from P. majori
from Samos and Akkaşdağı in having a relatively (compared
to the premolars) and absolutely longer upper molar row
(60.7–62.0 mm vs. 64.8–65.4 mm, based on the material
from Akkaşdağı), as well as in lacking a central fold (hypoconal spur) on P3 and P4, and an anterior cingulid on the
lower molars (Fig. 5; Kostopoulos 2009a). In addition, P. majori from Samos differs from the Şerefköy-2 material in the
presence of basal pillars on the upper molars and in having a
p4 bearing a centrally placed mesolingual conid and a fused
posterolingual conid and posterior stylid. By contrast, the
Şerefköy-2 specimens resemble P. pallasi from Samos and
Pikermi in all of these characters, as well as in the presence
of a posteriorly curved anterior style on P3, a more centrally
placed anterolabial cone on P4 (relative to P3), a strong mesostyle on the upper molars, a weak posterolabial conid on
p3 and p4, and a tendency for the mesolingual conid of p4 to
be oriented anteroposteriorly. P. pallasi from Pikermi differs,
however, in having distinct posterolingual and mesolingual
conids on p3, as well as distinct anterior and mesolingual
54
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
Length PM
140
130
120
110
100
70
72
80
78
76
74
Index P/M
Length pm
140
130
120
62
64
66
68
70
Index p/m
Palaeoryx pallasi
Şerefköy-2
Samos
Pikermi
Palaeoryx sp.
Kemiklitepe
Maragheh
Dytiko-2
Perivolaki
Palaeoryx
majori
Samos
Fig. 5. Scatter plot comparing the upper and lower premolar (P, p) / molar
(M, m) ratio against the complete tooth row length of Palaeoryx from several
localities.
conids on p4, although the degree of variability of these features within P. pallasi is not well known.
There is considerable size variation of both the upper
and lower tooth rows within Palaeoryx. The lower tooth row
from Şerefköy-2 (Lpm = 138.9 mm) appears close to that
from Maragheh (LPM = 139.3 mm). By contrast, the upper
tooth row (LPM = 111.5 mm) is considerably smaller than
both that from Maragheh (LPM = 140.8 mm) and a particularly large specimen from Pikermi (MNHN PIK2447; e.g.,
Kostopoulos and Bernor 2011), but close to P. pallasi from
Samos and other specimens from Pikermi. The available data
are insufficient for statistically sound conclusions; nevertheless, in the absence of marked morphological differences, we
suggest that the observed size variation may be intraspecific.
Stratigraphic and geographic range.—Upper Miocene; Balkans to Iran.
Genus Sporadotragus Kretzoi, 1968
Type species: Microtragus schafferi (Andree, 1926), Samos, Greece;
Late Miocene.
Sporadotragus parvidens (Gaudry, 1861)
Fig. 6.
Material.—MYŞE PV-2502, partial skull; MYŞE PV-1573,
frontlet preserving the basal part of the horn cores; MYŞE
PV-1300, partial left horn core; MYŞE PV-1522, right upper
tooth row with P2–M3; MYŞE PV-1412. right M3; MYŞE
PV-1423, left M3; MYŞE PV-1533, left M1; MYŞE PV-1511,
right mandibular body with p3–m3; MYŞE PV-2561, right
mandibular body with p2–m2; MYŞE PV-1429, right mandibular body with p4–m2; MYŞE PV-2556, right mandibular
body with m1–m3; MYŞE PV-1630, left mandibular body
with p3–m3; MYŞE PV-1407, PV-1574, left mandibular body
with p2–m2; MYŞE PV-2569, left mandibular body with p4–
m3; MYŞE PV-1406, PV-2559, left mandibular ramus with
p2–m3; MYŞE PV-1311, left mandibular ramus with p4–m2.
All from Şerefköy-2, Turkey, Late Turolian (Late Miocene).
Description.—This species is represented by at least 7 individuals. The frontals form a 105–115° angle along the sagittal
plane and appear moderately pneumatised above the orbits.
The supraorbital foramen (doubled on the left side in both
MYŞE PV-1573 and PV-2502) is small and round, placed
well below the pedicle, and opens directly into the orbit (Fig.
6A). The interfrontal suture is complex in outline and slightly
pinched between the horn cores (Fig 6A). The frontoparietal
suture is also complex, Y-shaped, and runs very close to the
base of the horn core. The postcornual fossa is shallow and
round. As preserved, the horn core is long (maximum length:
210 mm along the anterior surface) and gently curved posteriorly. It furthermore bears thin and discontinuous grooves
on its surface and shows no evidence of torsion (Fig. 6A).
In MYŞE PV-1300, a weak anterior keel runs along its basal
portion. The cross section of the horn core is elliptical at the
base, with its maximum transverse diameter located posteriorly (TD × 100/APD at the base = 75.3–76.4, n = 3; Table
3). The angle between the greatest anteroposterior diameter
of the horn core base and the sagittal plane ranges from 42
to 50°. Towards the tip, the cross section of the horn core
remains elliptical, but becomes symmetrical (TD × 100/APD
at 7 cm above the base = 72.9–78.3, n = 3; Table 3).
The upper premolars are moderately long compared to
the molars, with a premolar/molar ratio 69.7% (n = 1) (Fig.
6C, SOM 3: Table 1). P2 and P3 have a strong anterolabial
cone and bear a fossa unequally divided by a central fold
(Fig. 6B). The anterior style of P3 and P4 is also strong, but
the anterolabial cone of P4 is weak. The upper molars have
a strongly developed paracone, parastyle and mesostyle, a
weak metacone, and no basal pillars (entostyle) (Fig. 6B). A
metaconule fold is present on M2.
The lower premolar row is moderately short compared to
the molars, with a premolar/molar ratio of 58.3–62.4%, n = 2
Table 3. Horn core measurements (in mm) at the base and at 7 cm above
the base of Sporadotragus parvidens from Şerefköy-2 (Turkey). APD,
anteroposterior diameter; TD, transverse diameter.
Specimen
MYŞE PV-2502
MYŞE PV-1573
MYŞE PV-1300
TDbase
35.5
34.6
34.5
APDbase
44.3
46.0
45.1
TD7
28.1
27.8
29.0
APD7
35.9
35.6
39.8
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
5 cm
A
B
55
conid is well developed (Fig. 6C). Both the metastylid and
especially the entostylid of the lower molars are well marked
during early wear. There is no anterior cingulid. A low ectostylid is present on m1 (Fig. 6D).
Remarks.—Both the size and the overall horn core and dental
morphology of the material from Şerefköy-2 match those of
Sporadotragus, the taxonomic status of which was recently
revised by Geraads et al. (2006) and Kostopoulos (2009a).
The specimens from Şerefköy-2 have larger horn cores than
Sporadotragus vasili Geraads, Spassov, and Kovachev, 2006
from the SW Bulgarian localities of Kalimantsi and Strumyani
(Geraads et al. 2006, 2011; Fig. 7), which furthermore differs from the present material in having a fused interfrontal
suture, as well as a less curved horn core with a flat medial
and a wide anterior surface, strong longitudinal grooves, and
an anteromedial keel (Geraads et al. 2006). By contrast, the
specimens from Şerefköy-2 have several features in common
with Sporadotragus parvidens, mainly known from Pikermi, Samos (Greece) and Kemiklitepe-D (Turkey) (Bouvrain
1994; Kostopoulos 2009a). These include a small supraorbital foramen not located inside a pit, a complex and slightly
raised interfrontal suture, and a Y-shaped frontoparietal suture. In addition, both have a horn core that is long, sub-cylindrical and gently (but markedly) curved posteriorly, bears
irregular grooves, and shows no evidence of torsion or keels.
The flexion of the frontals along the sagittal plane appears
slightly stronger in the material from Şerefköy-2 (105–115°)
than in Sp. parvidens from Samos (100–105°); however, this,
as well as other minor differences (such as somewhat longer
51
C
47
5 cm
D
APDbase
(B–D)
43
39
Fig. 6. The bovid artiodactyl Sporadotragus parvidens (Gaudry, 1861) from
Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Frontlet (MYŞE PV2502) in anterior view. B. Right P2–M3 (MYŞE PV-1522) in occlusal view.
C. Right mandibular ramus with p3–m3 (MYŞE PV-1311) in lingual view.
D. Left mandibular ramus with p2–m3 (MYŞE PV-1406) in labial view.
(Fig. 6C, D, SOM 3: Table 2). The p3 and p4 have a strong
anterior stylid and a barely developed anterior conid (Fig.
6C). The mesolingual conid is simple and elongated on p3,
but rounded towards the base and slightly curved anteriorly
on p4. On both p3 and p4, the posterolingual conid fuses with
the posterior stylid during early wear and the posterolabial
35
30
35
TDbase
Sporadotragus parvidens Sporadotragus vasili
Strumyani
Şerefköy-2
Samos
Kalimantsi
Pikermi
25
40
Sporadotragus sp.
Kemiklitepe-D
Fig. 7. Scatter plot comparing the basal proportions of the horn cores of
Sporadotragus parvidens from Şerefköy-2 (Turkey), Samos and Pikermi
(Late Miocene, Greece) with Sporadotragus vasili from Kalimantsi and
Strumyani (Late Miocene, Bulgaria), and Sporadotragus from Kemiklitepe D (Late Miocene, Turkey). APDbase, anteroposterior diameter at the
base; TDbase, transverse diameter at the base.
56
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
horn cores in some specimens from Samos or the degree of
p4 molarisation) may well reflect intraspecific variation.
Stratigraphic and geographic range.—Upper Miocene; Aegean region.
Genus Skoufotragus Kostopoulos, 2009b
Type species: Pachytragus schlosseri (Andree, 1926), Samos, Greece;
Late Miocene.
Skoufotragus cf. Sk. schlosseri (Andree, 1926)
Figs. 8–10.
Material.—MYŞE PV-547, partial cranium with horn cores;
MYŞE PV-1571, 1576, female frontlet; MYŞE PV-1570, left
female horn core; MYŞE PV-1575, right horn core; MYŞE
PV-1579, partial left female horn core; MYŞE PV-2606, partial female horn core; MYŞE PV-1309, palate; MYŞE PV1512, PV-1516, PV-2570, right upper tooth row with P2–M3;
MYŞE PV-1513, PV-1514, PV-1515, right upper tooth row
with P3–M3; MYŞE PV-1622, right upper tooth row with
M1–M3; MYŞE PV-1491, right upper tooth row with M2–
M3; MYŞE PV-1523, right M3; MYŞE PV-1520, PV-1521,
left upper tooth row with M2–M3; MYŞE PV-1519, left upper tooth row with P2–M2; MYŞE PV-1525, left upper tooth
row with P4–M1; MYŞE PV-2571, left upper tooth row with
P3–M3; MYŞE PV-1410, left upper tooth row with M1–M3;
MYŞE PV-1517, left M2–M3; MYŞE PV-1312, left upper
tooth row with P2–P4; MYŞE PV-1315, left M3; MYŞE PV1434, PV-1436, PV-1313, M1 or M2; MYŞE PV-1532, left
P2; MYŞE PV-1510, PV-1542, PV-2560, right mandibular
body with p2–m3; MYŞE PV-1541, right mandibular body
with p3–m3; MYŞE PV-2551, right mandibular body with
p4–m2; MYŞE PV-2568, right mandibular body with m1–m3;
MYŞE PV-1546, left mandibular body with m2–m3; MYŞE
PV-1543, PV-1544, PV-2554, PV-2566, left mandibular ramus
with p2–m3; MYŞE PV-1540, left mandibular ramus with
p4–m3; MYŞE PV-2001, left mandibular ramus with p3–m1;
MYŞE PV-1156, left mandibular ramus with m2–m3. All from
Şerefköy-2, Turkey, Late Turolian (Late Miocene).
Description.—This is by far the most abundant bovid found
at Şerefköy-2 and represented by at least 10 individuals.
The opisthocranium is high, narrow, dolichocephalic (sensu
Bosscha-Erdbrink 1978) and has a straight dorsal profile in
lateral view (Fig. 8C, Table 4). The temporal lines are moderately developed and run parallel to each other posteriorly.
In posterior view, the occiput is triangular and bears a sharp
occipital crest (Fig. 8B, C) that ends dorsally in a strong
protuberance surrounded by deep scars. The nuchal crest is
well developed. The mastoid faces posterolaterally and the
paroccipital process is large and flattened. In lateral view,
the occipital condyles project posteroventrally, thus forming
a very acute angle with the occipital level. The basioccipital
is long, relatively narrow and bears a shallow, narrow longitudinal groove (Fig. 8D). The sharp and prominent (crestlike) posterior tuberosities of the basioccipital are oriented
perpendicular to the sagittal plane, whereas the weak anterior
tuberosities are oriented anteroposteriorly. The oval foramen
faces laterally. The preserved outline of the auditory bulla
indicates that it was large and bulbous.
The frontal contains large sinuses, one of which occupies
the pedicle and even reaches the base of the horn core. There
is no postcornual fossa. The horn core, inserted above the orbit, is sabre-like without keels or torsion (Figs. 8A, C, 9A). It
is moderately long, moderately curved posteriorly in lateral
view and strongly compressed mediolaterally along its entire
length (TD × 100/APD at the base: 40–57, n = 5; Table 4). In
cross section, the horn core forms an elongated ellipse, which
becomes narrower towards the tip. A deep furrow occasionally runs along the upper half of the posterior surface, and, in
combination with the strong mediolateral compression, may
give the impression of a distal posterior keel.
Two frontlets and three horn core specimens (MYŞE PV1570, PV-1571, PV-1576, PV-1579, and PV-2606; Fig. 9)
likely represent female individuals. The supraorbital foramen is large and round, and placed far anterior to the pedicle.
The interfrontal suture is open and simple in outline. The
horn core is thin and long (~140 mm) and inserted above
the back of the orbit. It is far removed from its counterpart,
weakly curved posteriorly and barely twisted homonymously (Fig. 9B). In cross section, the horn core is elliptical at the
base but becomes more compressed mediolaterally towards
the tip (Table 4). In anterior view, the divergence of the horn
cores is weak up to their mid-height, but stronger above.
The premolars are moderately long compared to the molars, with upper and lower premolar/molar ratios of 59.6–
65.6% (n = 5) and 60.3–66.6% (n = 4), respectively (Fig. 10,
Table 4. Cranial and horn core measurements (in mm) of Skoufotragus cf. Sk. schlosseri from Şerefköy-2 (Turkey). APD, anteroposterior diameter
at the base and at 10 cm above the base; Lfpo, sagittal length from the frontoparietal suture to the occipital protuberance; TD, transverse diameter
at the base and at 10 cm above the base; Watbc, width of the skull at the anterior tuberosities of the basioccipital; Wbrc, maximum width of the
braincase; Wptbc, width of the skull at the posterior tuberosities of the basioccipital. sin, left; dex, right.
Specimen
MYŞE PV-547 dex
MYŞE PV-547 sin
MYŞE PV-1571 dex
MYŞE PV-1575 dex
MYŞE PV-1570 sin
MYŞE PV-1576 dex
MYŞE PV-1576 sin
TDbase
29.29
32.2
34
33.36
25.92
22.42
23.16
APDbase
58.84
55.52
59.5
61.38
64.9
26.64
26.77
TD10
20.29
22.6
22.56
22.58
22.65
11.18
9.77
APD10
37.22
38.3
43.36
39.84
45.54
14.69
13.95
Lfpo
65.13
Wbrc
61.69
Wptbc
35.85
Watbc
19.94
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
A
57
C
B
5 cm
5 cm
(A, C)
D
5 cm
Fig. 8. The bovid artiodactyl Skoufotragus cf. Sk. schlosseri (Andree, 1926) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). Partial skull (MYŞE
PV-547) in anterior (A), posterior (B), lateral (C), and ventral (D) views.
SOM 3: Tables 1, 2). The upper molars have strong styles,
bear a fossetta (central islet) and lack entostyles (Fig. 10A,
B, D). P2 is bilobed lingually, whereas P3 has a trapezoidal
occlusal outline. Both have a strong anterolabial cone (Fig.
10A). The protocone of M1 protrudes lingually. The metastyle of M3 is strong and, in one specimen (MYŞE PV-1622),
flares distally (Fig. 10D). The mandibular body is shallow
and long (Fig. 10C). On the labial face, a second mental fora-
58
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
A
B1
B2
5 cm
5 cm
Fig. 9. The bovid artiodactyl Skoufotragus cf. Sk. schlosseri (Andree, 1926) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Frontlet (MYŞE
PV-1571) in anterior view. B. Female frontlet (MYŞE PV-1576) in anterior (B1) and lateral (B2) views.
A
B
5 cm
(A–D)
C
D
E
5 cm
(E, F)
F
Fig. 10. The bovid artiodactyl Skoufotragus cf. Sk. schlosseri (Andree, 1926) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Palate (MYŞE
PV-1309) in occlusal view. B. Left upper tooth row P2–M3 (MYŞE PV-1312) in buccal view. C. Right upper tooth row M1–M3 (MYŞE PV-1622) in occlusal view. D. Right mandibular ramus with p2–m3 (MYŞE PV-1510) in labial view. E. Left mandibular ramus with p2–m3 (MYŞE PV-1544) in occlusal
view. F. Right mandibular ramus with p2–m3 (MYŞE PV-1510) in occlusal view.
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
Remarks.—Protoryxoid bovids appear in the Eastern Mediterranean area as early as the Late Astaracian (e.g., Gentry
2000). Although they remained rare during the Vallesian,
they strongly radiated and dispersed during the Turolian,
especially in Anatolia and adjacent territories. They include
small to medium-sized antelopes of caprine/hippotragine
cranial appearance, but their taxonomy and evolutionary
relationships remain debated. Kostopoulos (2009a) partly
revised the Turolian protoryxoids from SE Europe assigned
to the so-called “Protoryx/Pachytragus complex” and recognised two distinct genera: Protoryx Major, 1891 and Skoufotragus Kostopoulos, 2009 (= partim Pachytragus Schlosser, 1904). The Şerefköy-2 specimens resemble Skoufotragus
in their dental morphology and in having (i) a narrow, long
braincase with a straight dorsal profile and parallel sides,
(ii) a triangular occiput and (iii) a sabre-like, mediolaterally compressed and uprightly inserted horn core (Kostopoulos 2009a: 364). Skoufotragus is known from the Turkish
Late Miocene assemblages of Kinik (Protoryx sp. of Köhler
1987), Kemiklitepe-A (Protoryx laticeps of Bouvrain 1994),
and Akkaşdağı (Pachytragus crassicornis of Kostopoulos
75
70
APD base
men appears below p3 (Fig. 10C). The p2 is simple without
an anterior conid, but with a strong mesolingual conid and
anterior stylid (Fig. 10E, F). The p3 has a well-developed
anterior conid and stylid, which become fused together with
wear. The mesolingual conid of p3 is oriented parallel to the
posterolingual conid, which in turn fuses with the weaker and
posterolingually directed posterior stylid during early wear
(Fig. 10E, F). The p4 resembles p3, but has an anteroposteriorly developed mesolingual conid (Fig. 10E). The lower molars have a strong meta- and mesostylids, but both elements
disappear with wear. There is no anterior cingulid. A basal
pillar appears on 5 out of 13 m1s, and 3 out of 13 m2s. The
third lobe of m3 is labially displaced and has an elongated,
semicircular occlusal outline; it bears a strong posterolingual
stylid on the upper half of the crown.
59
65
60
55
50
45
25
30
35
40
45
50
TD base
Skoufotragus laticeps
Maragheh
Samos
Skoufotragus
zemalisorum
Samos
Skoufotragus sp.
Kemiklitepe-A
Kinik
Skoufotragus schlosseri
Akkasdagi
Samos
Şerefköy-2
Protoryx carolinae
Pikermi
Fig. 11. Scatter plot comparing the basal horn core proportions of Skoufotragus from several sites and Protoryx from Pikermi (Late Miocene,
Greece). APDbase, anteroposterior diameter at the base; TDbase, trans-
verse diameter at the base.
2005), as well as from Samos, Greece and Maragheh, Iran
(Kostopoulos 2009a, Kostopoulos and Bernor 2011).
The Şerefköy-2 specimens differ from Skoufotragus laticeps from Samos, Kemiklitepe-A and Maragheh in having a
narrower braincase, as well as a shorter dorsal parietal sector, less developed anterior tuberosities of the basioccipital,
shorter, more slender, and more mediolaterally compressed
horn cores, and, on average, smaller tooth rows (but with a
similar premolar/molar ratio) (Figs. 11, 12). They are more
Component 2 (24%)
6
3
-20
-16
-12
-8
4
-4
8
12
-3
-6
Fig. 12. Principal component analysis of Skoufotragus from
Samos (Late Miocene, Greece), Akkaşdağı and Şerefköy-2
(both Late Miocene, Turkey), based on four cranial and horn
core variables (braincase width, dorsal length of parietal,
transverse diameter of the horn cores at the base, and anteroposterior diameter of the horn cores at the base) (modified
from Kostopoulos 2009a).
-9
-12
-15
Component 1 (62%)
Skoufotragus laticeps
Skoufotragus schlosseri
Skoufotragus zemalisorum
Akkasdagi
Serefkoy
16
60
similar to Sk. zemalisorum from Samos in terms of their
cranial proportions, but differ in having a narrower braincase (61.7 mm vs. 70–78 mm; Kostopoulos 2009a) and a
more anteroposteriorly expanded horn core (Figs. 11, 12).
Except for a somewhat longer lower premolar row (relative
to the molars), the Şerefköy-2 specimens closely resemble
Sk. schlosseri from Samos and Akkaşdağı in their size and
morphology (Figs. 11, 12), although the material from Samos
(Q5) is characterised by slightly longer and more divergent
horn cores bearing an anterior keel (the latter seems to be
more common in short-horned individuals; cf. Kostopoulos
2005: 777).
Female individuals of Skoufotragus are rare, but Gentry
(1971: 252, pl. 3: 3) interpreted AMNH 20687 as a female
individual of Sk. laticeps. The frontlet MYŞE PV-1576 from
Şerefköy-2 is very similar to this specimen, suggesting that
females of Sk. schlosseri were likely horned.
Genus Urmiatherium Rodler, 1889
Type species: Urmiatherium polaki Rodler, 1889, Maragheh, Iran; Late
Miocene.
Urmiatherium rugosifrons (Sickenberg, 1932)
Fig. 13.
Material.—MYŞE PV-1182, partial opisthocranium; MYŞE
PV-2503, frontlet; MYŞE PV-2504, frontlet of young individual; MYŞE PV-2599, axis+3rd cervical vertebra; MYŞE
PV-2506, axis; MYŞE PV-2505, 3rd cervical vertebra (possibly representing the same individual as MYŞE PV-2505).
All from Şerefköy-2, Turkey, Late Turolian (Late Miocene).
Description.—This bizarre species is represented by at least
two individuals and can unambiguously be identified based
on (i) its small size (Table 5); (ii) strong cranio-facial flexion;
(iii) strongly elevated, thick and pneumatised frontals; (iv) an
extremely shortened opisthocranium; (v) a reduced parietal
forming an obtuse angle with the occipital plane; (vi) an enlarged and thickened occipital with large occipital condyles
and a dorsally-facing occiput; (vii) a thickened basioccipital
with strong and completely fused posterior tuberosities, thus
forming an additional, oval facet for the atlas (Fig. 13C); and
(viii) thick, short, and moderately homonymously twisted
and grooved horn cores, originating very close from each
other and—in juveniles—bearing wide and well-defined lateral depressions (Fig. 13A, B).
Remarks.—Urmiatherium rugosifrons is the only representative of its genus from Asia Minor. Besides Şerefköy-2,
the species is only known from the neighbouring site of
Salihpaşalar (Kaya et al. 2012) and the Turolian of Samos
(e.g., Solounias 1981; Kostopoulos 2009a, 2014; Jafarzadeh et al. 2011; Table 5). Other possible occurrences of the
genus at Garkın, Kınık (Afyon) and Kayadibi (Konya) in
mid-western Anatolia (Alan 1997; Saraç 2003) still need to
be confirmed.
Stratigraphic and geographic range.—Upper Miocene; Asia
Minor.
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
Genus Sinotragus Bohin, 1935
Type species: Sinotragus wimani Bohlin, 1935, Locality 30, Shanxi
Province, North China.
cf. Sinotragus sp.
Fig. 14.
Material.—MYŞE PV-1409, partial left upper tooth row preserving only P2–P3; MYŞE PV-2553, left mandibular body
with p2–m3; MYŞE PV-2567, right mandibular body with
p2–m3. All from Şerefköy-2, Turkey, Late Turolian (Late
Miocene).
Description.—The only maxillary fragment preserves P2
and P3 (Fig. 14C, SOM 3: Table 1). P2 is weekly bilobed
lingually and has a trapezoidal occlusal outline, with a weak
anterior style, strong anterolabial cone, posterolabially protruding posterior style, and distolingually broadened posterolingual crista. P3 is subrectangular in occlusal view and bears
strong labial styles and ribs, as well as a weak lingual groove
dividing the distolingually expanded lingual cone (Fig. 14C).
A weak notch on the distal wall of the tooth emphasises the
posterior style. Traces of enamel on the distal occlusal surface suggest that the tooth originally had an additional distal
fossa or a strong central fold.
The lower premolar row is moderately short compared to
the molars, with premolar/molar ratios of 56.7% and 57.9%
(n = 2) (Fig. 14A, B, SOM 3: Table 2). The lower premolars
are narrow and short. The p2 lacks an anterior conid, but
bears an anteriorly located mesolingual conid and a week
anterior stylid. The mesolingual conid of p3 is elongated and
directed posteriorly. The posterolingual conid of p3 is strong.
The anterior conid and stylid are equally developed and fuse
together during advanced stages of wear. The anterior valley
is open. Unlike p3, p4 is distinctly molarised (Fig. 14A, B).
On the lingual wall, the anterior valley closes quickly during
wear and the mesolingual conid fuses with the posterior conid and stylid. Labially, there is a well-developed posterolabial conid. The lower molars bear a strong ectostylid and
have a gently undulating lingual wall covered by cement. The
Table 5. Cranial and horn core measurements (in mm) of Urmiatherium rugosifrons from Şerefköy-2 (Turkey) compared to Urmiatherium
rugosifrons from Samos (Greece) and Urmiatherium polaki from Maragheh (Iran). Data are from Kostopoulos (2009a) and Kostopoulos and
Bernor (2011). APDbase, anteroposterior diameter at the base; TDbase,
transverse diameter at the base; Wbi-co, bicondylar width of the skull;
Wbrc, maximum width of the braincase; Wptbc, width of the skull at
the posterior tuberosities of the basioccipital.
Specimen
Wbrc
MYŞE PV-1182
Wbi-co
57.6
Wptbc TDbase APDbase
28.1
MYŞE PV-2504
72.6
35.2
48.9
MYŞE PV-2503
92.7
~46.8
~70.8
42
78–130
Urmiatherium
rugosifrons, Samos
Urmiatherium
polaki, Maragheh
95–96
50–62
24–30
98
77–83
39
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
A
61
B
5 cm
C
D
Fig. 13. The bovid artiodactyl Urmiatherium rugosifrons (Sickenberg, 1932) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Juvenile frontlet
(MYŞE PV-2504) in anterior view. B. Frontlet (MYŞE PV-2503) in anterior view. C. Opisthocranium (MYŞE PV-1182) in ventral view. D. Axis + 3rd
cervical vertebra (MYŞE PV-2599) in dorsal view.
third lobe of m3 is relatively high and large, oval in occlusal
view, and bears a convex entoconulid.
Remarks.—The finely rippled enamel, advanced p4 morphology, reduced premolars, build-up of cement on the molars, large third lobe of the m3, strong basal pillars, and weak
development of the lingual stylids and ribs exclude this taxon
from the Late Miocene Antilopini sensu stricto. Together
with the inferred additional posterior fossa on P3, the distolingual widening of the lingual cone and the distolabial development of the posterior style on P2 and P3 are reminiscent
of the considerably larger Urmiatherium intermedium Bohlin, 1935, U. polaki Rodler, 1889, and Plesiaddax depereti
Bohlin, 1935. However, the Şerefköy-2 taxon differs from
the large-sized Late Miocene “ovibovines”, but resembles
the smaller U. rugosifrons from Samos (Kostopoulos 2009a:
371), in the development of the third lobe on m3, as well as
the weak buccolingual compression of the lower molars and
the degree of fusion of their lobes. U. rugosifrons and the
Şerefköy-2 species furthermore share a comparably reduced
premolar row. However, the former species is still about
30–35% larger and characterised by less derived lower premolars.
Although the available data are not sufficient for definite
conclusions, it is worth mentioning that a right upper tooth
row from Muğla (MNHN TRQ 974) resembles the material
from Şerefköy-2 in its size and premolar morphology. The
molars of TQR 974 considerably differ from those of Urmiatherium and Plesiaddax, but resemble those of Sinotragus
wimani Bohlin, 1935 from China (Bohlin 1935: pl. 16: 8, 9).
Unfortunately, the lower dentition of Si. wimani is unknown,
preventing a direct comparison with Şerefköy-2. TRQ 974
is accompanied by a frontlet from the same region (MNHN
TRQ 973), which, based on its morphology and size, likely represents a female Si. occidentalis Geraads, Güleç, and
Kaya, 2002. This species is well-documented in Muğla by
62
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
Karadenizli et al. 2005) and characterises the Final Bovid
Assemblage of Samos (~7.0–6.8 Ma; Kostopoulos 2009a;
Koufos et al. 2009b). The co-occurrence of U. rugosifrons
and Skoufotragus cf. Sk. schlosseri at Şerefköy-2 thus places
the latter close to the Middle–Late Turolian boundary (~6.8
Ma; e.g., Steininger 1999; Agusti et al. 2001).
A1
A2
Palaeobiogeographical and
palaeoenvironmental analysis
5 cm
B
C
Fig. 14. The bovid artiodactyl cf. Sinotragus from Şerefköy-2 (Turkey),
Late Turolian (Late Miocene). A. Left mandibular ramus (MYŞE PV2553) in labial (A1) and lingual (A2) views. B. Right mandibular body
(MYŞE PV-2567) in occlusal view. C. Left upper P2–P3 (MYŞE PV-1409)
in occlusal view.
cranial, but not dental, elements (Geraads et al. 2002). Given
the particular dental characters of TRQ 974, the similar size
of TQR 974 and TRQ 973, and their common geographic
provenance, we speculate that both specimens belong to Si.
occidentalis. In addition, we tentatively refer the Şerefköy-2
specimens to Sinotragus based on their similar dental morphology.
Biochronology of Şerefköy-2
Kaya et al. (2012) suggested a Middle Turolian age for the
Şerefköy-2 assemblage and stressed its structural similarity
to the mammalian palaeocommunity from Samos. The results of the present study help to refine this biochronological
assessment. Among the bovid recovered from Şerefköy-2,
Gazella capricornis and Palaeoryx pallasi are the most
widespread, ranging from the Balkans to Iran and the northern coast of the Black Sea. Their co-existence at Şerefköy-2
is strongly indicative of a Middle–Late Turolian age for
this faunal assemblage, as is the presence of Sporadotragus
parvidens. The latter is so far only known from Samos and
Pikermi (Greece), although the genus occurs from the southern Balkans all the way to Afghanistan and possibly China
and Mongolia (e.g., Geraads et al. 2006). Urmiatherium rugosifrons certainly occurs at Mytilinii-1A and most likely
also at Q5 of Samos (Kostopoulos 2009a), indicating an age
younger than 7.1 Ma (Koufos et al. 2009b). By contrast,
Skoufotragus schlosseri is present at Akkaşdağı (<7.1 Ma;
Protoryxoid bovids represent more than 60% (more than
80% together with gazelles) of the Şerefköy-2 bovid community in terms of both the minimum number of individuals
(n = 27) and the number of identifiable specimens (n = 102).
In sharp contrast, boselaphines and spiral-horned antelopes,
both of which are common in contemporaneous faunas of the
sub-Paratethyan province, are absent from Şerefköy-2. Even
though taphonomic and sampling biases may have exaggerated these results, the absence of these taxa indicates that
they were at least rare in the local mammal community. On
the other hand, the presence of Urmiatherium rugosifrons, so
far only known from neighbouring Samos, and of a western
Asian representative of Sinotragus, known mainly from the
surroundings of Şerefköy (i.e., the Muğla-Yatağan Basin;
e.g., Geraads et al. 2002), increase the local character of the
Şerefköy-2 mammal assemblage.
In our genus-level correspondence analysis, the first
three components (out of 15) account for 23.2%, 20.8%, and
14.1% of the inertia, respectively (Fig. 15A). Both in the first
(axes 1 and 2; Fig. 15A) and the second (axes 1 and 3; not
shown) plane, the continental Greek Turolian bovid associations are sharply distinguished in taxonomic composition
and relative abundances from most Anatolian bovid assemblages (including Samos), whereas the Turkish assemblages
of Mahmutgazi, Sivas and Akkaşdağı, and the Iranian fauna
of Maragheh, occupy an intermediate position, rather close
to the Greek bovid faunas (Fig. 15A). The primary polarizing taxa include Paraoioceros, Plesiaddax, Protragelaphus,
Nisidorcas, Majoreas, and Urmiatherium. There is a clear,
time-controlled taxonomic shift in the Greek bovid assemblages from Nikiti-2 (Early Turolian) to Dytiko (Late Turolian), with an increase of the magnitude of Miotragocerus,
Gazella, and Protragelaphus against Nisidorcas, and Tragoportax (Fig. 15A).
In the east, the situation seems much more complicated
than in continental Greece. The Middle Turolian bovid assemblages of Asia Minor, Mytilinii-1A, B, C, Mytilinii-3, Kemiklitepe-A, B, Kinik, and Şerefköy-2 form a distinct cluster,
characterised by Skoufotragus, Sporadotragus, Palaeoryx,
Urmiatherium, and Paraoioceros. The older (Early Turolian)
but geographically close assemblages of Kemiklitepe-D and
Mahmutgazi are separated from this group mainly because
of the high percentages of Criotherium, Majoreas, Oioceros,
and Plesiaddax. The central-eastern Anatolian bovid faunas
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
A
63
16
2.0
15
KTD
1.5
2.0
1.5
Cc
Axis 2
1.0
1
NIK
0.5
0.0
14
PXM
2
RZO PER
MAR 13
34
Sivas
-0.5
5
Mahm
PIK DYTI
6 7
Akkas
12
8
MTLA
9
Seref
KTAB
MYT
10 11
Kinik
-1.0
-1.5
-2.0
-2.0
-1.5
-1.0
0.0
-0.5
0.5
RZO
B
1.0
2.5
Axis 1
Cc
NIK
DYTI
Akkas
Sivas
PXM
MAR
PER
PIK
Kinik
MYT
Mahm
Seref
KTD
MTLABC
KTAB
Fig. 15. Correspondence analysis of Turolian (Late Miocene) bovid associations from the sub-Paratethyan zoogeographic province, based on relative taxonomic abundances at the genus level (A) and relative size abundances at each site (B). See text for explanation, SOM 2 for details of the methodology, and
SOM 1: Table 1 for the dataset. Analysed variables: 1, Nisidorcas; 2, Tragoportax; 3, Prostrepsiceros; 4, Palaeoreas; 5, Protragelaphus; 6, Miotragocerus;
7, Gazella; 8, Urmiatherium; 9, Palaeoryx; 10, Paraoioceros; 11, Skoufotragus/Protoryx; 12, Sporadotragus; 13, Plesiaddax; 14, Oioceros; 15, Majoreas;
16, Criotherium. Squares represent Early Turolian assemblages; circles represent Middle–Late Turolian assemblages. In the spectra of relative size abundances (pie charts) grey represents small (<50 kg), white medium, and black large-sized bovids (>150 kg). Abbreviations: Akkas, Akkaşdağı; Cc, Çorak-yerler; DYTI, Dytiko-1, 2, 3; KTAB, Kemiklitepe A, B; KTD, Kemiklitepe D; Mahm, Mahmutgazi; MAR, Maragheh (mainly levels MMTT7-13); MTLABC,
Mytilinii-1A, B, C; MYT, Mytilinii-3; NIK, Nikiti-2; PER, Perivolaki; PIK, Pikermi; PXM, Prochoma; RZO, Ravin de Zouaves-5; Seref, Şerefköy-2.
exhibit a similar pattern, with the Early Turolian Çorak-yerler assemblage being separated from those of Sivas and Akkaşdağı (late Early and late Middle Turolian, respectively).
The bovid assemblages of Çorak-yerler and Kemiklitepe-D
are much more alike in their taxonomic structure than are
the roughly contemporaneous faunas of continental Greece
(Nikiti-2 and Ravin de Zouanes-5) (Fig. 15A). By contrast,
Sivas and, to a lesser degree, Akkaşdağı, more closely re-
semble assemblages from continental Greece to the west and
Maragheh to the east than contemporary faunas from Asia
Minor.
In the correspondence analysis using broader taxonomic bins, the first two components (out of four) account for
51.8%, and 25.2% of the inertia, respectively (Fig. 16A). The
results are similar to those of the genus-level analysis and
show no major changes between the Early and mid-Late Tur-
64
ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015
A
B
2.4
RZO
PER
PIK
KTD
KTAB
MYT
DYTI
OVIB
2
KTD
Axis 2
1.6
MTLABC
1.2
0.8
Seref
Sivas
0.4
MAR
0
-0.4
SPIRA
NIK
RZO
Akkas
Mahm
Cc
PER DYTI
PXM BOSE
PIK
Sivas
GAZE
Seref
MTLABCPROTO
Akkas
KTAB
MYT
Kinik
-0.8
-0.9
-0.6
-0.3
0
0.3
Axis 1
0.6
0.9
1.2
1.5
1.8
Fig. 16. Correspondence analysis of Turolian (Late Miocene) bovid associations from the sub-Paratethyan zoogeographic province based on taxonomic
bins (A) and diet spectra of several local bovid associations based on dental wear patterns (B). See text for explanation, SOM 2 for details of the methodology, and SOM 1: Table 2 for the dataset. Abbreviations: Akkas, Akkaşdağı; Cc, Çorak-yerler; DYTI, Dytiko-1, 2, 3; KTAB, Kemiklitepe A, B; KTD,
Kemiklitepe D; Mahm, Mahmutgazi; MAR, Maragheh (mainly levels MMTT7-13); MTLABC, Mytilinii-1A, B, C; MYT, Mytilinii-3; NIK, Nikiti-2;
PER, Perivolaki; PIK, Pikermi; PXM, Prochoma; RZO, Ravin de Zouaves-5; Seref, Şerefköy-2. Dashed-line represents a convex hull of Early Turolian
bovid assemblages; BOSE, GAZE, PROTO, OVIB, SPIRA, represent analysed variables (SOM 1: Table 2). Pie charts represent the spectra of feeding
preferences: black, browsers; grey, mixed feeders; white, grazers.
olian assemblages from continental Greece, all of which are
dominated by spiral-horned antelopes and boselaphines (Fig.
16A). The same taxa fare less well in contemporary Anatolia,
where they and “ovibovines” are increasingly replaced by
gazelles and protoryxoids. This trend is much sharper in the
western part of the region; in central-eastern Anatolia, assemlages instead remain more similar to those of continental
Greece and Iran (Fig. 16A).
Although based on a much more limited dataset lacking
Turolian faunas from Greece, Bibi, and Güleç (2008: fig. 11)
found a similar taxonomic trend in the Anatolian bovid assemblages, including the separation of Çorak-yerler and Kemiklitepe-D from the remaining assemblages. However, they
also identified a cluster comprising Kemiklitepe-A, B and
Sivas, despite their marked taxonomic differences (Tragoportax constitutes 41.5% at Sivas but is absent from KTAB;
“Pachytragus/Protoryx” constitutes 3.8% at Sivas vs. 39.6%
at KTAB; Bibi and Güleç 2008), size distribution and diet
spectra (see below). This grouping thus seems rather improbable and probably stems from the inclusion of the Greek
Vallesian assemblage of Nikiti-1. This may have resulted in
the compression of the remaining faunas within multispace,
as in de Bonis et al. (1994: fig. 4).
Faunal body size distributions generally support the results of the CFA (Fig. 15B). Thus, the Turolian faunas of
continental Greece contain very few large taxa (<4%) and,
with the exception of the mostly small (<50 kg) species constituting the earliest Turolian assemblage of Nikiti-2, show a
balance between small and medium-sized forms (Fig. 15B).
A similar pattern occurs in central-eastern Anatolia (Fig.
15B), although the dataset is much more restricted here. By
contrast, large and small-sized bovids are balanced and predominate at Kemiklitepe-D, whereas from Mahmutgazi to
Şerefköy-2 large taxa decline and medium-sized bovids increase considerably, with percentages reaching 80% in some
cases (Fig. 15B).
The diet spectra of the assemblages compared here do
not follow the regional pattern revealed by the CFA and
size distribution analyses. Instead, they point to a general
rise of mixed feeders at the expense of grazers in younger associations, as shown by comparisons of, for instance,
KTD with KTAB and Şerefköy-2 in western Anatolia, Sivas
with Akkaşdağı in eastern Anatolia, and Ravin de Zouaves-5
with Perivolaki, Pikermi, and Dytiko in Greece (Fig. 16B).
In addition, several roughly contemporaneous faunas show
similar diet spectra despite marked differences in their geographical location, taxonomic composition and size distribution (e.g., Nikiti-2 vs. Çorak-yerler, Dytiko vs. Akkaşdağı,
and Pikermi vs. Kemiklitepe-A, B and Mytilinii-1A, B, C;
Figs 15A, B, 16B).
KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY
65
Discussion and conclusions
Acknowledgements
As in the study by Bibi and Güleç (2008: 516), there is
no straightforward correlation between the component axes
arising from the taxonomy-based CFA and environmental
trends, at least at a generalised Greco-Anatolian level. Instead, the age and geographical location of the sites, two
parameters not directly involved in the analysis, seem to
control much of the distribution of the data. Thus, in line
with preliminary suggestions by Kostopoulos and Bernor
(2011), our analysis revealed a “Greek” cluster and a “SW
Anatolian” cluster, whereas the eastern Anatolian Turolian
bovid assemblages possibly form a third group more closely
related to Maragheh in Iran.
Like in continental Greece, the general increase of mixed
feeders relative to grazers throughout the Turolian in eastern
Anatolia is not accompanied by a major reorganization of the
structure of the bovid community, with the exception of an
increase in the number of protoryxoids. Instead, size ratios
generally remain stable, with small taxa always contributing
more than 50% (based on the number of identifiable specimens). During approximately the same time period, small
taxa slightly decrease in number (by about 10%) in SW Anatolia, whereas medium-sized taxa considerably increase (by
about 30%). Protoryxoids at least quadruple at the expense of
boselaphines and “ovibovines”, while the relative abundances of gazelles and spiral-horned antelopes remain stable at a
moderate and low level, respectively.
These results generally agree with ungulate regional endemicity along the Greco-Iranian longitudinal axis, as suggested by Costeur (2009: fig. 4), and are fully compatible
with the conclusions of de Bonis et al. (1994). The partial isolation of SW Anatolia from continental Greece was
thoroughly discussed by Kostopoulos (2009b, and literature
therein) and likely resulted from both the mid-Tortonian/
Maeotian transgression into the Aegean domain and the latest
Miocene invasion of the Paratethyan Sea. By contrast, the
partial faunal isolation of SW Anatolia from adjacent regions
to the east may be related to the broad Mio-Pliocene Central
Anatolian lake system stretching between the Pontides in
the north and the Taurides-Isparta Angle barrier in the south
(e.g., Görür et al. 1995; Veen et al. 2009; Alçiçek 2010; Mehmet Cihat Alçiçek personal communication 2012).
Our study demonstrates that similar ecological signals
need not necessarily imply identical size distributions or taxonomic content, even for roughly contemporaneous, neighbouring faunas. This implies that the faunal composition of
herbivores at various points of a supposedly homogeneous
biogeographic province may depend more on historical (i.e.,
phylogeographic) relationships and limitations than the type
of vegetation cover. Assuming that the diet spectrum of a local herbivore assemblage directly reflects vegetational and/
or climatic conditions, it therefore seems that local bovid
communities adjust to the available habitat, rather than (or
prior to) being reorganised by migratory movements and
replacements.
We thank Tanju Kaya and Serdar Mayda (both Ege University, Izmir,
Turkey), Robert Scott (Rutgers University, New Brunswick, New
Jersey, USA), and Gildas Merceron (Poitiers University, France) for
their help and support in the field and lab, and Mehmet Cihat Alçiçek
(Pamukkale University, Denizli, Turkey) for helpful discussions on
SW Anatolia geodynamics. Thanks are also due to Kaye Reed (Arizona State University, Tempe, USA), Denis Geraads (CNRS MNHN,
Paris, France) and Felix G. Marx (University of Otago, New Zealand)
for their fruitful comments and suggestions, as well as to Katerina Vasileiadou (Natural History Museum of the Petrified Forest of Lesvos,
Greece) for linguistic improvements. This work was supported by a
“Wenner-Gren International Collaborative Research Grant” (“Environmental Dynamics of Western Eurasian Hominids during the Late
Miocene”; PI: Robert Scott), as well as grants from Ege University
(TTM/001/2008 and TTM/001/2010) and the Scientific and Technological Research Council of Turkey (108Y047).
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Veen, J.H. ten, Boulton, S.J., and Alçiçek, M.C. 2009. From palaeotectonics to neotectonics in the Neotethys realm: The importance of kinematic decoupling and inherited structural grain in SW Anatolia (Turkey).
Tectonophysics 473: 261–281.
Vislobokova, I. 2005. The importance of Late Miocene faunal exchanges
between Eastern Mediterranean areas and Central Europe. Annales de
Paléontologie 91: 241–255.
Western, D. 1979. Size, life history and ecology in mammals. African
Journal of Ecology 17: 185–204.
http://app.pan.pl/SOM/app60-Kostopoulos_Karakutuk_SOM.pdf
SUPPLEMENTARY ONLINE MATERIAL FOR
Dimitris S. Kostopoulos and Seval Karakütük
Late Miocene bovids from Şerefköy-2, SW Turkey, and their position
within the sub-Paratethyan biogeographic province
Published in Acta Palaeontologica Polonica 2015 60 (X): xxx-xxx.
http://dx.doi.org/10.4202/app.2012.0019
SOM 1
Table 1. Relative abundances of bovid genera from Greek, Turkish, and Iranian Turolian
assemblages.
Table 2. Proportions of size, taxonomic, and feeding categories of bovid assemblages from
Greek, Turkish and Iranian Turolian assemblages.
SOM 2
Correspondence analysis, size and diet spectra.
References
SOM 3
Table 1. Upper tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi,
Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, and cf. Sinotragus from Şerefköy-2
(Turkey).
Table 2. Lower tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi,
Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, and cf. Sinotragus from Şerefköy-2
(Turkey).
3.00
70.83
39.6
6.81
68.2
1.16
4.11
41.72
43.14
22.17
7.50
12.50
4.55
2.62
5.96
17.65
1.25
4.17
12.50
2.09
3.49
1.58
8.61
6.86
3.30
2.02
2.50
1.00
42.86
41.51
33.75
57.53
4.00
23.00
50.00
28.57
4.00
16.98
16.25
15.07
1.25
1.37
5.00
1.25
15.07
4.17
8.33
1.05
6.82
10.17
3.16
6.62
35.08
18.90
8.86
0.66
3.30
34.41
2.36
2.02
4.11
1.99
5.88
4.65
32.91
21.47
2.27
3.20
21.84
15.12
0.95
21.47
7.85
0.95
1.74
2.85
23.58
9.72
21.86
Gazella
Protragelaphus
66.67
10.66
3.00
Paraoioceros
7.58
2.54
Majoreas
Prostrepsiceros
Oioceros
Plesiaddax
22.84
14.29
3.77
3.75
Criotherium
Urmiatherium
19.70
54.82
Nisidorcas
3.00
Tragoportax
Miotragocerus
Palaeoryx
1.52
Palaeoreas
NIK
RZO
Cc
KTD
Sivas
Mahm
PXM
MYT
KTAB
PER
Kinik
PIK
MAR7-13
MTLABC
Seref
Akkas
DYTI
Sporadotragus
Skoufotragus/
Protoryx
SOM 1. Table 1. Relative abundances of bovid genera from Greek, Turkish, and Iranian Turolian assemblages.
4.55
9.14
10.00
14.29
30.19
27.50
10.96
8.33
39.58
3.66
18.18
38.95
10.44
34.44
23.53
45.28
19.43
SOM 1. Table 2. Proportions of size, taxonomic, and feeding categories of bovid assemblages from Greek, Turkish, and Iranian Turolian
assemblages.
NIK
RZO
Cc
KTD
Sivas
Mahm
PXM
MYT
KTAB
PER
Kinik
PIK
MAR
MTLABC
Seref
Akkas
DYTI
PROTO
1,5
0
3
14,3
3,8
12,5
0
87,5
53
8,9
72,7
7,3
5,7
56,3
67,6
25,5
2
BOSE
19,7
54,8
2
0
41,5
36,2
57,5
0
0
36,1
6,8
29,1
14,2
7,3
2,9
5,7
36,5
OVIB
0
0
5
43
0
16,3
0
0
0
0
0
0
4,1
2
5,9
0
0
SPIRA
74,2
36
80
28,6
17
7,5
31,5
4,2
8,3
51,3
2,3
24,7
59,5
0
0
23,6
42,1
GAZE
4,6
9,1
10
14,3
37,7
27,5
11
8,3
39,6
3,7
18,2
39
16,5
34,5
23,5
45,3
19,4
Small
78,8
45,2
86
42,8
54,7
33,8
42,5
12,5
39,6
47,2
20,4
61,9
73,1
34,4
23,5
68,9
39,7
Medium
19,7
54,8
9
14,3
45,3
48,7
57,5
83,3
47,9
50,8
79,6
34,6
19
57
69,6
27,8
58,3
Large
1,5
0
5
42,9
0
17,5
0
4,2
12,5
2
3,5
7,9
8,6
6,9
3,3
2
Br
Mix
Gr
35,6
?
42,9
17
0
?
0
11,3
64,4
?
57,1
71,7
87,5
20,8
30,6
4,2
39,6
45,3
8,3
39,6
24,1
21,8
76,5
1,7
30,5
24,5
6,6
6
48,3
43,1
50
47,8
21,2
32,4
43,4
46,2
NISP
66
197
100
56
53
80
73
24
48
191
44
344
316
151
102
212
247
Abbreviations.—Greek faunas: NIK, Nikiti-2; RZO, Ravin de Zouaves-5; PXM, Prochoma; MYT, Mytilinii-3; PER, Perivolaki; PIK, Pikermi;
MTLABC, Mytilinii-1A,B,C; DYTI, Dytiko-1,2,3. Turkish faunas: Cc, Çorak-yerler; KTD, Kemiklitepe D; Sivas, Sivas; Mahm, Mahmutgazi;
KTAB, Kemiklitepe A, B; Kinik, Kinik; Seref, Şerefköy-2; Akkas, Akkaşdağı. Iranian faunas: MAR, Maragheh (mainly levels MMTT7-13).
NISP, number of identified specimens. Size categories: small, <50 kg; medium, 50–150 kg; large, > 150 kg. Diet categories: Br, browsers; Mix,
intermediate feeders; Gr, grazers. See SOM 2 for further details.
SOM 2. Correspondence analysis, size and diet spectra
We performed two separate correspondence analyses. The first followed the basic
concepts of Bibi and Güleç (2008: 515, 519), based on the relative abundances of
bovid genera per site for Turolian (Late Miocene; 8.7–5.0 Ma; Steininger 1999)
assemblages from Greece (n = 8), Turkey (n = 8) and Iran (n = 1). Relative
abundances of bovid genera were estimated based on the number of identified
specimens (NISP), which is closely related to the minimum number of individuals
(MNI) [a regression analysis of MNI-NISP pairs of 60 bovid genera from 10 of the
Greek and Turkish faunas resulted r2 = 0.93 and p (uncorr) = 2.8E - 33; analysis not
shown, but available on request], and weakly related to the number of bovid genera
per site [r2 = 0.49; p (uncorr) = 0.001]. Counts for Sivas and Çorak-yerler are from
Bibi and Güleç (2008), for Kinik and Mahmutgazi from Köhler (1987), for
Akkaşdağı, Perivolaki, Mytilinii-3, and Mytilinii-1A,B,C from Kostopoulos (2005,
2006, 2009b), for Nikiti-2 from Kostopoulos and Koufos (1999) and personal data,
for Maragheh from Kostopoulos and Bernor (2011), and for Kemiklitepe D and A,B
from Bouvrain (1994). Data for Pikermi include specimens from the NHML, MNHN,
and KNUA collections. Counts for Ravin de Zouaves-5, Prochoma, and Dytiko are
based on personally collected data. Single occurrences at the genus level were
excluded from the analysis.
In contrast to Bibi and Güleҫ (2008), we grouped bovid genera into five
taxonomic units in our second correspondence analysis: PROTO, including
protoryxoid bovids (Skoufotragus/Pachytragus, Protoryx, Sporadotragus, and
Palaeoryx);
BOSE,
including
Late
Miocene
boselaphines
(Miotragocerus,
Tragoportax, and Samokeros); GAZE, including Late Miocene representatives of
Gazella and its allies; OVIB, including Late Miocene “ovibovines” (Criotherium,
Urmiatherium, Sinotragus, and Plesiaddax); and SPIRA, including Late Miocene
spiral-horned antelopes (Oioceros, Paraoioceros, Dytikodorcas, Hispanodorcas,
Prostrepsiceros, Palaeoreas, Majoreas, Nisidorcas, Protragelaphus, and Pheraios).
These taxonomic bins, based principally on phenetic similarity (but in some cases also
phylogenetic relationships), simply reflect the basic taxonomic structure of the bovid
assemblages and allowed us to (1) emphasize general patterns in the taxonomic
structure; (2) incorporate genera with low abundances that otherwise require
exclusion from the analysis; and (3) overcome differing taxonomic opinions (e.g.,
Bibi and Güleç 2008).
We do not follow Bibi and Güleç (2008) in assuming that all species of a
genus tend to have similar environmental responses, but agree with their assumption
that the absolute abundance of a taxon or group of taxa through time is itself timeindependent. As body size appears to be a crucial ecological parameter both at the
level of the organism and the community level (e.g., Western 1979; Eisenberg 1990),
we assume that the size structure of a bovid assemblage is important in understanding
its paleoecological context and paleogeographic relationships. We therefore also
performed an analysis of the relative abundances of small (less than 50 kg), medium,
and large bovids (more than 150 kg) per site. All size categories include taxa of
different tribal affiliations, thus making them sufficiently independent from a
taxonomic point of view.
Finally, we classified all of bovid genera into three main groups of feeding
preferences (based on their relative abundances), so as to be able to show the diet
spectra of several local bovid associations. These groups consisted of grazers
(including intermediate grazers), mixed feeders, and browsers (including intermediate
browsers). However, because a single genus may include species with rather distinct
feeding behaviors (e.g., Prostrepsiceros; see Merceron et al. 2004, 2005a; Solounias
et al. 2010), bovid genera from Ravin de Zouaves 5 (RZO), Perivolaki (PER), Pikermi
(PIK), Dytiko (DYTI), Mytiliniii-3 (MYT) and Mytilinii-1A,B,C (MTLABC) were
classified strictly based on the results of the dental-wear pattern analyses (including
both microwear and mesowear) conducted by Koufos et al. (2006, 2009), Merceron
et al. (2005a, b), and Solounias et al. (2010). This means that the same genus may
appear as a grazer in one bovid association, but as a browser in another, depending on
the local species record (e.g., Merceron et al. 2005a; Solounias et al. 2010). When
available, we also considered differences within the same taxonomic unit, as in
Tragopotrax rugosifrons from RZO and PER (Merceron et al. 2005a: 480; Koufos et
al. 2006: 207), and Gazella from MTLAB (including three species with different
ecological profiles; Koufos et al. 2009b). Owing to the absence of similar studies on
Turkish faunas, we classified the bovids from Sivas, Kemiklitepe D and A-B,
Şerefköy-2, and Akkaşdağı by assuming their feeding preferences to resemble those
of taxa from other sites close in time and geography. We did not analyse the
ecological spectra of Maragheh, Kinik, Corak-yerler, Mahmutgazi, and Nikiti-2,
either because of a lack of data or because of other, similar studies on these sites
currently in progress.
References
Bibi, F. and Güleç, E.S. 2008. Bovidae (Mammalia: Artiodactyla) from the late
Miocene of Sivas, Turkey. Journal of Vertebrate Paleontology 28: 501–519.
Bouvrain, G. 1994. Les gisements de mammifères du Miocène supérieur de
Kemiklitepe, Turquie: 9. Bovidae. Bulletin du Muséum National d’Histoire
Naturelle 4, sect. C. 16 (1): 175–209.
Eisenberg, J.F. 1990. The behavioral/ecological significance of body size in the
Mammalia. In: J. Damuth and B. MacFadden (eds), Body Size in Mammalian
Paleobiology Estimation and Biological Implications, 25–37. Cambridge
University Press, Cambridge.
Köhler, M. 1987. Boviden des Türkischen Miozäns (Kanozoikum und Braunkohlen
der Türkei). Paleontologia i Evolucio 28: 133–246.
Koufos, G.D., Merceron, G., Kostopoulos, D.S., Vlachou, T., and Sylvestrou, I. 2006.
The late Miocene vertebrate locality of Perivolaki, Thessaly, Greece. 11.
Palaeoecology and Palaeobiogeography. Palaeontographica Abteilung A 276:
201–221.
Koufos, G.D., Kostopoulos, D.S., and Vlachou, T. 2009. The Late Miocene Mammal
Faunas of the Mytilinii Basin, Samos Island, Greece: New collection. 16.
Chronology. Beiträge zur Paläontologie 31: 397–408.
Kostopoulos, D.S. 2005. The Bovidae (Artiodactyla, Mammalia) from the Late
Miocene
mammal
locality
of
Akkaşdağı
(Central
Anatolia,
Turkey).
Geodiversitas 27: 747–791
Kostopoulos, D.S. 2006. The late Miocene vertebrate locality of Perivolaki, Thessaly,
Greece. 9. Cervidae and Bovidae. Palaeontographica Abteilung A 276: 151–183.
Kostopoulos, D.S. 2009b. The Pikermian Event: temporal and spatial resolution of
the Turolian large mammal fauna in SE Europe. Palaeogeography,
Palaeoclimatology, Palaeoecology 274: 82–95.
Kostopoulos, D.S. and Bernor, R.L. 2011. The Maragheh bovids (Mammalia,
Artiodactyla):
systematic
revision
interpretation. Geodiversitas 33: 649–708
and
biostratigraphic-zoogeographic
Kostopoulos, D.S. and Koufos, G.D. 1999. The Bovidae (Mammalia, Artiodactyla) of
the Nikiti-2 [NIK] faunal assemblage (Chalkidiki peninsula, N. Greece). Annales
de Paléontologie 85: 193–218.
Merceron, G., Blondel, C., Brunet, M., Sen, S., Solounias, N., Viriot, L., and Heintz,
E. 2004. The late Miocene paleoenvironment of Afganistan inferred from dental
microwear in artiodactyls. Palaeogeography, Palaeoclimatology, Palaeoecology
207: 143–163.
Merceron, G., Bonis, L. de, Viriot, L., and Blondel, C. 2005a. Dental microwear of
fossil bovids from northern Greece: paleoenvironmental conditions in the eastern
Mediterranean during the Messinian. Palaeogeography, Palaeoclimatology,
Palaeoecology 217: 173–185.
Merceron, G., Bonis, L. de, Viriot, L., and Blondel, C. 2005b. Dental microwear of
the late Miocene bovids of northern Greece: Vallesian/Turolian environmental
changes and disappearance of Ouranopithecus macedoniansis? Bulletin de la
Societé géologique de France 176: 475–484.
Solounias, N., Rivals, F., and Semprebon, G.M. 2010. Dietary interpretation and
paleoecology of herbivores from Pikermi and Samos (late Miocene of Greece).
Paleobiology 36: 113–136.
Steininger, F. 1999. Chronostratigraphy, geochronology and biochronology of the
Miocene “European land mammal mega-zones” (ELMMZ) and the Miocene
“mammal-zones” (MN-zones). In: G.E. Rössner, K. Heissig, and V. Fahlbusch,
(eds.), The Miocene Land Mammals of Europe, 9–24. Verlag Friedrich Pfeil,
Munich.
Western, D. 1979. Size, life history and ecology in mammals. African Journal of
Ecology 17: 185–204.
SOM 3
SOM 3. Table 1. Upper tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf.
Sk. schlosseri, and cf. Sinotragus from Şerefköy-2 (Turkey). L, length; W, width; sin, left; dex, right.
Specimen
LPM
Gazella cf. capricornis
MYŞE PV-2575
MYŞE PV-2572sin 48.9
MYŞE PV-2572dex 48.0
Palaeoryx pallasi
MYŞE PV-2573
111.5
MYŞE PV-1295
MYŞE PV-1293
MYŞE PV-1294
Sporadotragus parvidens
MYŞE PV-1533
MYŞE PV-1412
MYŞE PV-1423
MYŞE PV-1522
69.7
Skoufotragus cf. Sk. schlosseri
MYŞE PV-1519
MYŞE PV-1513
MYŞE PV-1512
96.3
MYŞE PV-1516
86.3
MYŞE PV-1514
MYŞE PV-1520
MYŞE PV-1515
MYŞE PV-1521
MYŞE PV 1309 dex 90.2
MYŞE PV 1309 sin 88.9
MYŞE PV-1312
MYŞE PV-1525
MYŞE PV-2570
86.2
MYŞE PV-2571
MYŞE PV-1410
MYŞE PV-1491
MYŞE PV-1622
MYŞE PV-1532
MYŞE PV-1312
MYŞE PV-1517
MYŞE PV-1315
MYŞE PV-1523
MYŞE PV-1434
MYŞE PV-1436
MYŞE PV-1313
?Sinotragus sp.
MYŞE PV-1409
LP
LM
LP2
WP2
LP3
WP3
LP4
WP4
21.5
20.5
20.4
28.7
28.6
7.7
7.6
6.9
6.6
5.7
5.5
7.3
7.1
6.8
6.8
6.3
6.1
6.4
6.5
6.7
7.3
7.7
6.9
46.8
65.4
15.1
13.0
16.8
16.7
18.0
18.1
15.2
14.7
15.1
16.9
17.0
17.6
64.8
LM1 WM1
43.6
32.0
35.6
33.7
57.2
59.7
54.2
59.0
WM2 LM3 WM3
7.6
7.4
9.9
9.2
10.0
10.5
9.5
9.6
11.0
10.8
8.9
9.2
20.3
20.4
22.9
19.8
24.8
21.4
22.4
21.8
26.7
27.1
25.3
24.7
26.8
26.4
15.7
18.8
14.7
12.2
11.3
9.7
12.7
13.6
14.8
12.2
12.8
14.0
12.2
14.9
14.7
14.5
30.5
LM2
12.4
9.9
7.7
9.8
7.9
9.7
8.5
14.8
12.2
16.0
11.4
9.3
9.1
9.8
10.1
11.8
9.9
8.8
11.05
9.6
10.6
12.2
11.1
11.9
12.6
12.4
11.2
13.2
11.9
12.0
11.7
11.4
7.9
9.8
11.4
8.2
15.1
18.3
19.0
16.6
18.9
17.4
16.2
14.8
17.7
13.9
18.7
20.7
20.32
16.4
15.4
15.2
12.3
11.0
10.4
12.2
16.5
16.8
16.7
17.6
21.2
21.7
18.4
21.3
19.6
19.9
14.1
14.3
16.0
15.1
16.0
16.5
20.9
22.8
20.0
21.7
21.5
18.6
20.6
20.8
19.7
18.8
17.5
15.3
20.0
14.7
14.5
17.3
14.6
17.8
16.7
19.8
18.6
21.0
21.1
20.2
14.3
18.3
14.0
14.8
16.1
20.4
19.6
19.1
20.0
23.0
11.8
11.6
12.8
13.2
12.5
18.5
15.0
22.1
21.0
22.2
11.7
13.7
13.7
19.9
19.5
14.3
14.7
15.0
52.0
36.6
35.7
55.8
55.3
12.4
11.5
9.3
9.85
10.5
8.8
12.8
12.5
11.3
11.4
11.4
12.0
12.4
11.7
10.6
11.0
9.5
34.8
53.9
9.9
6.8
11.5
13.0
8.9
11.6
11.8
11.5
8.1
13.2
10.2
9.1
8.8
8.2
54.0
8.5
8.4
11.2
8.9
7.4
9.1
11.6
8.1
SOM 3. Table 2. Lower tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf.
Sk. schlosseri, and cf. Sinotragus from Şerefköy-2 (Turkey). L, length; W, width.
Specimen
Lpm
Gazella cf. capricornis
MYŞE PV-2562
54.8
MYŞE PV-2557
54.6
MYŞE PV-2563
54.7
MYŞE PV-2558
55.0
MYŞE PV-2564
52.6
MYŞE PV-2565
MYŞE PV-2000
56.7
MYŞE PV-1528
Palaeoryx pallasi
MYŞE PV-2574
138.9
MYŞE PV-1599
137.0
MYŞE PV-2552
Sporadotragus parvidens
MYŞE PV-1574
MYŞE PV-2561
MYŞE PV-1511
MYŞE PV-1311
MYŞE PV-2556
MYŞE PV-1630
MYŞE PV-1407
MYŞE PV-1406
72.6
MYŞE PV-1429
MYŞE PV-2569
MYŞE PV-2559
Skoufotragus cf. Sk. schlosseri
MYŞE PV-1543
90.0
MYŞE PV-1540
MYŞE PV-1541
MYŞE PV-1510
90.0
MYŞE PV-1542
101.2
MYŞE PV-1156
MYŞE PV-2001
MYŞE PV-1546
MYŞE PV-2566
MYŞE PV-2560
MYŞE PV-2568
MYŞE PV-2554
92.3
MYŞE PV-2551
?Sinotragus sp.
MYŞE PV-2553
65.3
MYŞE PV-2567
65.7
Lp
Lm
Lp2
Wp2
Lp3
Wp3
Lp4
Wp4
Lm1
Wm1
Lm2
Wm2
Lm3
Wm3
20.3
20.1
20.8
20.2
18.7
5.1
5.3
4.8
5.7
5.0
3.0
3.1
2.8
3.1
3.1
2.9
3.8
4.1
3.8
4.0
3.8
3.8
4.1
8.9
7.5
7.6
7.6
8.9
4.7
4.2
4.8
4.8
4.9
4.6
4.4
8.8
9.1
8.7
8.2
8.5
8.1
9.3
6.0
6.0
5.9
5.5
7.6
6.9
7.2
7.2
6.7
6.9
7.8
8.7
21.5
35.7
34.4
34.9
34.4
34.6
32.0
35.7
11.4
10.5
10.5
10.8
10.6
9.9
11.5
6.3
6.3
6.1
6.6
5.9
6.4
6.1
15.9
14.9
16.0
15.4
15.3
14.1
15.5
15.9
5.4
6.2
5.3
6.6
5.7
6.2
5.8
6.1
54.0
55.1
84.0
82.7
14.2
13.7
7.9
8.6
18.9
18.6
10.7
11.9
20.7
21.4
11.6
14.8
22.2
18.7
12.2
27.0
25.9
23.4
13.5
18.5
33.7
37.7
29.6
13.5
16.4
6.8
6.7
4.4
3.8
10.4
10.5
9.9
6.3
5.3
6.4
10.6
11.0
10.9
11.2
6.5
6.4
6.5
6.5
12.2
13.1
12.6
12.0
11.1
7.3
7.9
7.8
8.3
7.3
8.2
7.1
8.4
7.9
17.4
7.1
5.4
6.3
6.3
19.5
19.5
8.0
8.3
13.2
11.1
11.9
8.5
9.4
9.6
9.6
10.3
19.5
9.2
11.4
7.8
8.8
9.6
9.5
8.6
14.6
15.3
13.8
14.8
14.3
15.2
14.8
13.6
16.6
16.1
14.5
20.8
22.9
9.1
9.8
9.3
10.1
11.5
9.9
17.8
18.9
19.3
17.6
19.6
18.3
11.2
9.0
9.9
11.2
10.3
10.2
24.9
26.0
25.9
24.5
27.4
26.3
10.6
8.7
10.3
10.3
9.9
9.5
18.0
17.5
18.4
17.6
16.8
17.4
10.0
10.6
11.7
12.3
12.5
25.3
25.1
10.3
10.1
10.7
11.3
11.3
12.7
13.3
9.7
9.2
19.9
18.6
9.0
8.7
27.3
28.5
43.9
43.5
27.0
27.8
44.5
6.8
8.0
4.2
4.6
10.5
10.0
10.0
27.5
47.0
47.2
7.0
4.0
10.2
5.2
10.1
5.8
11.4
6.6
34.8
35.0
38.5
56.0
61.8
63.8
56.2
63.8
35.1
35.2
9.9
10.5
5.6
5.9
12.1
13.2
6.2
6.7
12.0
6.0
5.8
5.7
10.8
10.9
11.2
11.2
12.2
7.1
7.1
7.2
7.1
7.0
13.6
12.9
8.1
7.4
14.2
14.0
7.6
8.0
16.2
16.9
13.3
15.9
7.5
16.9
9.5
10.0
10.4
5.3
5.3
11.6
11.9
6.5
7.0
13.4
13.4
8.1
8.2
15.7
15.0
10.3
12.2
37.1
55.6
55.6
11.1
5.9
13.0
7.0
13.7
13.7
8.6
8.0
13.7
13.0
11.2
10.5
24.3
24.0
42.0
42.3
7.0
7.5
4.9
4.8
8.5
8.3
5.8
5.8
8.8
8.5
6.4
9.4
8.9