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Teeth of early generations of Early Pleistocene

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Sinyaya Balka/Bogatyri site (Sea...

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DOI: 10.1016/j.quaint.2015.08.007

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 Quaternary International 420 (2016) 306e318

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Teeth of early generations of Early Pleistocene elephants (Mammalia,

Elephantidae) from Sinyaya Balka/Bogatyri site (Sea of Azov Region,
Russia)
Vera S. Baigusheva a, Vadim V. Titov b, *, Irina V. Foronova c
a
Azov Museum-Reserve, Moskovskaya str., 38/40, Azov, Russia
b
Institute of Arid Zones SSC RAS, Southern Scientic Centre of the Russian Academy of Science, Chekhov str., 41, Rostov-on-Don, 344006, Russia
c
V.S. Sobolev Institute of Geology and Mineralogy, Siberian Branch of the Russian Academy of Science, pr. Koptiuga, 3, Novosibirsk, 630090, Russia

a r t i c l e i n f o a b s t r a c t

Article history: Based on study of a series of early generations' teeth DP4/dp4 e M1/m1 of elephants from the Late
Available online 23 October 2015 VillafranchianeEarly Galerian locality Sinyaya Balka/Bogatyri (Taman peninsula, Russia) the variability of
their characteristics is established. There is no clear normal distribution of main diagnostic dental pa-
Keywords: rameters in the sample. This conrms the assumption of the possible presence of two elephant taxa in
Early Pleistocene the structure of Taman Faunistic complex from south Eastern Europe, which was made earlier on the
Taman Faunistic complex
basis of the M3 teeth. The main part of the collection is represented by remains of the late meridionaloid
Archidiskodon
elephant subspecies Archidiskodon meridionalis tamanensis, typical for late Early Pleistocene of Eastern
Dental characteristics
Variability
Europe and neighboring regions. Individual specimens of the sample may refer to other form of ele-
phants with similar morphological teeth characteristics, probably to Phanogoroloxodon mammonthoides.
Though the lesser diagnosticity of early generation teeth in comparison with the M3/m3, it was found
that they can be used when determining the taxonomic forms of elephants, even for the meridionaloid
elephant subspecies.
2015 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction
A determination of systematic position of fossil elephants is
mainly based on features of skull structure and morphometric data
of molars of last generation. Craniological features they specify the
taxa of relatively high rank, while evolutionary changes within the
Abbreviations: AMZ, Azov historical, archeological and paleontological
museum-reserve, Azov, Russia; GGM, State geological museum of V.I. Vernadskiy,
Moscow, Russia; GIN, Geological institute of Russian Academy of Sciences, Moscow,
Russia; IPG, Institute of paleobiology, Tbilisi, Georgia; KM, Krasnodar state historical
and archeological museum-reserve of E.D. Felicyn, Krasnodar, Russia; NMC, Don
Cossacks' history museum, Novocherkassk, Russia; PIN, Paleontological museum of
Russian Academy of Sciences, Moscow, Russia; PKM, Pyatigorsk regional museum,
Pyatigorsk, Russia; PRKM, Primorsk regional museum, Primorsk, Ukraine; ROMK,
Rostov-on-Don Regional museum, Rostov-on-Don, Russia; SMZ, Stavropol state
museum-reserve, Stavropol, Russia; SSC, Southern Scientic centre of Russian
Academy of Sciences, Rostov-on-Don, Russia; TM, Taganrog regional museum,
Taganrog, Russia; ZIN, Zoological Institute of Russian Academy of Sciences, St.
Petersburg, Russia.
* Corresponding author.
E-mail addresses: vvtitov@yandex.ru (V.V. Titov), irina_foronova@mail.ru
(I.V. Foronova).
genera and species can be traced by the teeth structure. The pres-
ervation of fossil elephants' teeth and their study have established
systematic and phylogenetic positions of these leading fossils of
Pleistocene continental deposits.
Almost all the features of elephants' teeth structure can be
evaluated in numerical values, that change consecutively both in
the ontogeny and phylogeny (Garutt and Foronova, 1976). The tooth
crown size, number of plates, their length, and enamel thickness
increase in each successive generation in ontogenesis. The number
of enamel plates and lamellar frequency increase, but the length of
single plates and enamel thickness decreased from archaic to
advanced forms in phylogeny. These morphometric characteristics
used for taxonomic diagnosis of Proboscidea are most clearly
expressed on teeth of the two last generations (M2 and M3). Based
on the study of teeth M3/m3 variability and skulls of southern
elephant Archidiskodon meridionalis s.l. in early Pleistocene of south
Eastern Europe, three taxa are dened: A. m. gromovi (Middle Vil-
lafranchian Khapry Faunal complex); A. m. meridionalis (Late Vil-
lafranchian Psekups Faunal complex); and A. m. tamanensis (Late
VillafranchianeEarly Galerian Taman Faunal complex) (Gromov,
1948; Dubrovo, 1964; Garutt and Alexeeva, 1964; Alexeeva and

http://dx.doi.org/10.1016/j.quaint.2015.08.007
1040-6182/ 2015 Elsevier Ltd and INQUA. All rights reserved.
 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318
Garutt, 1965; Alexeeva, 1977; Garutt, 1977, 1986; Garutt et al., 1977;
Baigusheva, 2000; Baigusheva and Titov, 2001, 2010, 2012; Kahlke
et al., 2011).
There are disagreements regarding the genus afliation and the
species of meridionaloid elephants. Some researchers do not
recognize the validity of the genus Archidiskodon and include
southern elephants in the genus Mammuthus (Maglio, 1973;
Aguirre and Morales, 1990; Lister, 1993, 1996; Lister and Sher,
2001; Lister et al., 2005; Palombo and Ferretti, 2005; Essen van,
2009, and others). According to another point of view, the genus
Archidiskodon is autonomous and is regarded as a primitive stage of
elephants of tribe Mammuthini Brookes (Osborn, 1942; Dubrovo,
1960, 1964, 2001; Baigusheva, 1971; Alexeeva, 1977; Azzaroli,
1977; Garutt, 1986, 1998; Foronova and Zudin, 1999; Baigusheva
and Titov, 2012, and others). Some of these paleontologists rate
them as mammuthoid elephants. The present authors hold the
opinion that Late Pliocene e Early Pleistocene mammuthoid ele-
phants should be attributed to the separate genus Archidiskodon,
being a paraphyletic group, according to a non-cladistic attitude
(c.f. Lister, 1996). This point of view is based on distinct differences
between Late PlioceneeEarly Pleistocene and MiddleeLate Pleis-
tocene representatives of this elephants' tribe.
Diagnosis of early generation teeth of elephants is complex due
to the fact that the characteristics are not expressed clearly, and are
less stable because of the transgression of parameters, as well as
due to insufcient study. In this context, the teeth DP2/dp2 e M1/
m1, especially single nds, practically are not used for taxonomic
denitions. However, the information about the structure and
variability of early generation teeth belonging to different members
of the mammoth lineage is of scientic interest. Some information
about them can be found in the literature.
Among the works dedicated to analysis of dental system of
mammoth lineage's elephants and morphometric characteristics of
teeth of all generations, including premolars, are publications of
Garutt and co-authors (Garutt and Foronova, 1976; Garutt, 1977;
Garutt and Bajgu
seva, 1981). In these papers, dental morpho-
metric characteristics are presented in tables and graphs, and show
the variability of characteristics, including teeth of early genera-
tions of different taxa. On the basis of large serial material,
Fig. 1. Some Eastern European Middle and Late Villafranchian localities with Archidiskodon meridionalis remains: 1 e Sinyaya Balka/Bogatyri, 2 e Tsimbal, 3 e Sarkel, 4 e Port-Katon,
5 e Semibalki, 6 e Samarskoe, 7 e Nogaisk, 8 e Akhalkalaki, 9 e Old Georgievsk sand pit, 10 e Saratovskaya/Psekups, 11 e Linentsovka, 12 e Khapry, 13 e Mokriy Chaltir, 14 e
Kobiakova Balka, 15 e Kosiakino, 16 e Kryzhanovka, 17 e Kotlovina.
307
including the localities discussed in this article, Garutt and
Foronova (1976) proposed a method that uses not extreme values
of transgressing digital characteristics, but optimal (mode) ones,
which are the most frequent for this form. However, in that pub-
lication the numerical characteristics of A. meridionalis' teeth were
given as a whole, without subdivision into chronological sub-
species: A. m. meridionalis (Nesti, 1825) and A. m. tamanensis
Dubrovo, 1964. This makes it difcult to use it in this study.
In papers which study elephants' teeth from Pleistocene of the
North Sea area (Essen van and Mol, 1996; Essen van, 2009), there is
a comparison of characteristics of several teeth generations of
Mammuthus meridionalis, Mammuthus trogontherii, and
M. primigenius from some localities. However, these authors give
the parameters of teeth DP3/dp3 e M1/m1 only in graphs, which
mainly explain the changing of general size and hypsodonty index
of teeth. When analyzing M. trogontherii teeth from Sssenborn
(Germany), Guenther (1969) compared parameters of teeth of
trogontherii mammoth with those of meridionaloid elephants and
woolly mammoths. He also showed the variability of teeth,
including mm2 ( DP3/dp3), mm3 ( DP4/dp4) and M1/m1 in the
graphs. Maglio (1973) gives a summary table with teeth measure-
ments (including DP2/dp2-M1/m1) of meridionaloid elephants
from multiple Italian Villafranchian sites.
There is also a number of publications describing single nds of
Archidiskodon and Mammuthus teeth of early generations from Italy
(Falconer and Cautley, 1846; Friant, 1959), Germany (Untermabfeld;
Dubrovo, 2001), France (Saint-Vallier; Gue rin, 2004); Georgia
(Akhalkalaki; Vekua, 1962), as well from Russia: Azov Sea, Lower
Don and the Taman Peninsula (Gromov, 1977; Maschenko, 2002;
Baigusheva and Titov, 2012), Stavropol Region (Gabunia, 1961),
Trans-Baikal (Udunga; Kalmykov and Maschenko, 2006), and
Yakutia (Vilyuisk; Dubrovo, 1953). Dubrovo (2001) gives charac-
teristics of individual teeth DP4/dp4 of elephants and mammoths
from some localities from Ukraine and Russia, including the Taman
Peninsula. The early generation teeth of M. primigenius are studied
and described in more detail (e.g. Maschenko, 2002).
There has been no detailed description of representative series
of southern elephant teeth of early generations until today.
Therefore, the teeth collection of the later form of southern
308 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318

elephant A. m. tamanensis from Eastern Europe from site Sinyaya
Balka/Bogatyri (Taman Peninsula, Russia) is important. It allows us
to reveal the characteristic morphological features and the vari-
ability of teeth DP4/dp4 and M1/m1 parameters of the form under
study. Data on individual teeth DP2/dp2 and DP3/dp3 (Table 1) are
also provided. It is possible to use the result when deciding issues of
onto- and phylogeny of the mammoth lineage's elephants.
mammonthoides which lived together with southern elephants
during the late Early Pleistocene in Eastern Europe. Ph. mammon-
thoides Garutt, 1958 has similar teeth morphology, but differs
markedly from Archidiskodon in skull characteristics (Figs. 2 and 3;
Garutt, 1958, 1986, 1995). According to the taxon's author, the
phanagorian elephant was a member of the tribe Phanagorox-
odontini, which he put at the base of the divergence of the lines

Table 1
Tooth measurements of teeth DP3, dp3, DP2 of Archidiskodon meridionalis from some Villafranchian localities of southern East Europe and western Asia.

DP3, dp3, DP2 Length Width Hight Dental formula (number Lamellar frequency Length of Enamel thickness Hypsodonty Wear
(mm) (mm) (mm) of plates and talons) (at 5 cm) single plate (mm) index (H/W) stage

Late VillafranchianeEarly Galerian

Tsimbal, Sinaya Balka/Bogatyri (Taman peninsula, Russia)
dp3, KM 1802/2 P-6, sin 53.8 33.2 e e5t e e 2.2 e 5
dp3, ZIN 25994/15, dex 81.4 49.0 e t6t 3.25 12.4 1.05 e 5
dp3, ZIN 25994/28, sin >60 43.0 e e5t 5.0 9.48 1.0 e 5
dp3, AMZ 30814/1, sin 70.1 33.4 t5t 4.75 10.57 1.4 e 5
Samarskoe (Sea of Azov Region, Russia)
dp2, ROMK 1247, sin 23.0 20.0 12 t3t 7.5 e 1.0 0.6 0
dp3, ROMK 1247, sin >61 40.0 33 5e 5.0 e 1.8 0.825 1
Akhalkalaki (Georgia)
DP3, IPG 20, sin 72.0 42.0 >46 t7t 4.5 e 0.9 e 5
Late Villafranchian
Georgievsk (Stavropol Region, Russia)
DP3, SMZ no >65 35.0 e ~6t 5.0 e 1.0 e 5
Middle Villafranchian
Linentsovka, Khapry (Sea of Azov Region, Russia)
DP2, GIN 300/13, dex 25.8 20.4 21.0 t4t e 4.4 e 1.03 1
DP2, GIN 300/122, dex 31.0 29.2 20.0 t3t 6.0 e 1.0 e 1
DP2, GIN 300/125 32.0 22.0 18.0 t4t e 6.2 e 0.82 1
dp3, ROMK L-53, sin 73.6 29.2 39.0 t6t 4.5 10.7 1.1 1.34 3
dp3, ROMK L-1487, dex 73.8 41.6 e t7t 4.9 10.3 1.2 e 4
dp3, GIN 300/124, sin 59.6 31.2 29.6 t6t 5.5 8.8 1.1 0.95 4
Kobiakova Balka (Sea of Azov Region, Russia)
DP3, ZIN 29071, dex >59.5 34.8 30.4 t6t 5.0 8.8 0.8 0.87 4
Kuruksay (Tadzhikistan)
DP3, GIN 3848/67 >52 34.8 39.6 e5t 4.5 9.4 e 1.14 5

1.1. Site

Site Sinyaya Balka/Bogatyri is situated on the northern shore of

the Taman Peninsula close to Cape Bogatyri near the village Za
Rodinu (Temryuk district, Russia) (Fig. 1). The geological age of the
locality is traditionally dened approximately as late Early Pleis-
tocene, 1.2e0.9 Ma (Vangengejm et al., 1991) based on the evolu-
tionary level of fauna. According to new data, including
archaeological ones, it is dated ca. 1.5e1.2 Ma (Shchelinsky et al.,
2010). The typical mammal assemblage of this Late Villa-
franchianeEarly Biharian European mammal age includes Trogon-
therium cuvieri Fischer von Waldheim, Castor tamanensis
Verestchagin, Canis tamanensis Verestchagin, Archidiskodon mer-
idionalis tamanensis, Elasmotherium caucasicum Borissiak, Equus cf.
major Boule, Cervidae gen., Bison sp., and Tragelaphini gen.
(Verestchagin, 1957; Baigusheva et al., 2008; Sotnikova and Titov,
2009; Shchelinsky et al., 2010). Based on the fauna from this lo-
cality, the Taman faunal complex was identied. It was typical for
the end of the Early Pleistocene (Eopleistocene) of south Eastern
Europe (Gromov, 1948).
There are different views about the specic structure of ele-
phants from Sinyaya Balka/Bogatyri. Belyaeva (1925) allocated two
forms: Elephas meridionalis and E. trogontherii. Verestchagin
(1957) also noted several species: E. meridionalis, E. trogon-
therii, and E. antiquus. Dubrovo (1964) described new subspecies
A. meridionalis tamanensis on the basis of numerous materials from
the locality. She considered all morphological differences as indi-
Fig. 2. Upper right tooth M1 no. KM-40 of Phanogoroloxodon mammonthoides from
vidual variability of a single form. Garutt (1958) ascribed one tooth Northern Caucasus, holotype (Early Pleistocene, Psekups locality; by Garutt, 1958): a, b
from the Sinyaya Balka collection to Phanogoroloxodon e occlusal surface (1  1). Collection of Krasnodar museum-reserve.
 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318
Fig. 3. Skull no. KM-40 of Phanogoroloxodon mammonthoides from Northern Caucasus,
holotype (Early Pleistocene, Psekups locality; by Garutt, 1958): a e dorsal view, b e
lateral view. Collection of Krasnodar museum-reserve.
Fig. 4. Stratigraphical position of some Eastern European Early Pleistocene localities of meridionaloid elephants.
309
Mammuthini and Elephantini. Lister et al. (2005) assumed a pres-
ence of remains of progressive M. meridionalis and early
M. trogontherii in the collection, reected in the bimodal charac-
teristics of the teeth. Our research of last and penultimate molar
generations of 2e4 stages of wear conrmed the bimodal distri-
bution of M3/m3 and m2 features (number of plates, lamellar fre-
quency, and index of hypsodonty). In the Sinyaya Balka/Bogatyri
sample, there are morphologically similar teeth, which can be
divided into 2 groups (Baigusheva and Titov, 2012). The rst group
is reliably correlated with A. m. tamanensis. The taxonomic status of
the second form (Elephantidae form 2) is still debatable. The last
molars of A. m. cf. tamanensis which were described from the
similar age Sarkel locality (lower Don River, southern European
Russia, late Early Pleistocene), containing the remains of Taman
theriofauna representatives, show a bimodal distribution of fea-
tures on diagrams by the enamel thickness and lamellar frequency
(Nikolskiy et al., 2014).
310 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318
2. Materials and methods
We studied a series of teeth of later forms of the southern
elephant Archidiskodon meridionalis tamanensis, from the site of
Sinyaya Balka/Bogatyri. There are characteristics of 15 specimens of
DP4, 8 dp4, 18 M1 and 24 m1 were involved in the present study
(Table 6). The bone remains from this locality were collected at
different times by different researchers. In 1912, studies were
conducted by the Russian academician Gubkin. In 1957, these were
supplemented by the expedition of the Paleontological Institute
RAS (Moscow) under the guidance of Dubrovo. Separate specimens
from the 1990 excavations under the leadership of Zhegallo are
stored in the Vernadsky Geological Museum (Moscow). New nds
were collected during the excavations of the site Sinyaya Balka/
Bogatyri led by Shchelinsky and Kulakov (Institute for the History of
Material Culture of Russian Academy of Sciences) in 2004e2014.
They are stored in the Azov Museum-Reserve (Azov). The article
uses the data from other localities (Baigusheva and Titov, 2008) of
similar geological age from the Sea of Azov Region and Ciscaucasia
(Figs. 1 and 4): Tsymbal (Taman Peninsula), Sarkel (Lower Don),
Fig. 5. Upper teeth of Archidiskodon meridionalis tamanensis, locality Sinyaya Balka/
Bogatyri, Taman peninsula, Russia, Late VillafranchianeEarly Biharian: a e DP4 PIN
1249/165, left; b e DP4 PIN 1249/179, right; e M1 AMZ 30814/12, left, 6 wearing
degree; d e M1 PIN 1249/285, left, 4 wearing degree; Phanogoroloxodon aut Archi-
diskodon: e  M1 PIN 1249/006, left, 4 wearing degree.
Sredniy Egorlyk, Samarskoe, Semibalki, and Port Katon, Nogaisk
(Northeastern Sea of Azov Region).
To determine variability and possibility of using the elephant
teeth of young generations for biostratigraphic purposes, we
compared a sample of teeth from Sinyaya Balka (65 specimens)
with a series of Archidiskodon meridionalis gromovi Garutt, Alex-
eeva, 1965 (43 specimens) from Northeast Middle Villafranchian
localities from the Sea of Azov Region (Liventsovka, Khapry,
Kobyakovskaya Balka, Mokriy Chaltyr') from the vicinity of Rostov-
on-Don (Baigusheva, 1971; Garutt and Bajgu
seva, 1981; Titov,
2008). This collection was generally gathered by one of the au-
thors (V.S. Baigusheva), and is stored in the funds of the Rostov
Regional Museum of Local History (Rostov-on-Don) and Azov
Museum-Reserve (Azov). Separate ndings are present in collec-
tions of the Zoological Institute (St. Petersburg) and the Geological
Institute of Russian Academy of Sciences (Moscow).
Fig. 6. Lower teeth of Archidiskodon meridionalis tamanensis, locality Sinyaya Balka/
Bogatyri, Taman peninsula, Russia, Late VillafranchianeEarly Biharian: a e dp3 AMZ
30814/1, left, 5 wearing degree; b e dp4 AMZ 30814/2, right, 4 wearing degree; e m1
30426/37, right, 5 wearing degree; d e m1 Z 29899/2, right, 4 wearing degree.
 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318 311

Parameters of some teeth of A. meridionalis meridionalis from
Late Villafranchian localities Saratovskaya/Psekups (Krasnodar
Region, Russia) and Georgieskiy sand pit (Stavropol Region,
Russia) are also used. A comparison with the tooth M1 of Ph.
mammonthoides holotype, described from Early Pleistocene site
Psekups from Ciscaucasia, was carried out (Garutt, 1958, 1992,
1995).
To study premolars DP2/dp2 e DP4/dp4 and molars M1/m1 the
same morphometric approaches and methods as for the teeth of
last generation, M3, were used. They are traditionally used in the
study of Proboscidean remains (Gromova, 1964, 1965; Dubrovo,
1960, 1964; Aguirre, 1969; Guenther, 1969; Maglio, 1973; Garutt
and Foronova, 1976; Foronova and Zudin, 1986, 1999). Crown di-
mensions (length, width, height) and lamellar frequency, average
length of a single plate, enamel thickness, and type of the incipient
wear gures on the occlusal surface of trinomial plates of tooth
were analyzed.
Lamellar frequency is the number of plates on 100 mm or
50 mm of crown. The index is an average based on measurements
taken in the middle part of the lateral and medial surfaces of a
crown. The measure of average length of single plate was calcu-
lated by taking the average of several values of the quotient from
the division by 5 of the ve plate's length (included inter-plate
spaces), measured in different parts of the crown at lingual and
labial sides.
Average enamel thickness was calculated using an average from
the possible number of measurements taken within the entire
chewing surface. Thus, statistical reliability was achieved.
The wear gures on the occlusal surface of trinomial plates of
antiquoid ( d ), meridionaloid (d  d) or intermediate (d d d)
types were noted. Due to considerable wear of the majority of teeth
(87% of teeth are of the 4th and 5th wear stage) the hypsodonty
index was not considered in this study. We consider that averages
of various parameters of the teeth are essential for the sample. At a
rather large variability of characteristics they allow to dene a
common evolutionary level of the investigated taxa (Gromova,
1965). Particular attention was paid to teeth parameters' changes
depending on the wearing degree. The resulting digital data are
presented in tables (Tables 1e6).

Table 2
Tooth measurements of upper teeth DP4 of Archidiskodon meridionalis from some Villafranchian localities of southern East Europe and western Asia.

DP4 Length Width Hight Dental formula (number Lamellar frequency Length of Enamel thickness Hypsodonty Wear
(mm) (mm) (mm) of plates and talons) (at 10 cm) single plate (mm) index (H/W) stage

Late VillafranchianeEarly Galerian

Sinaya Balka/Bogatyri (Taman peninsula, Russia)
PIN 1249/83, sin >71 59.2 >65.4 5t 7.5 13.4 1.5 1.10 5
PIN 1249/149, dex >110 56.2 88.4 t8t 6.5 14.5 2.2 1.57 2
PIN 1249/165, sin 128.0 58.0 >69.0 t9t 8.0 11.8 2.1 1.19 4
PIN 1249/179, dex 126.0 61.0 >66.0 t8t 7.25 13.4 1.85 1.08 5
PIN 1249/189, dex >131 66 107 e8t 7.0 13.2 1.4 1.62 3
PIN 1249/190, sin 134.0 74.2 e t8t 6.5 13.9 1.9 e 4
PIN 1249/199, dex e 50.4 >63.2 5 7.5 11.9 1.6 1.25 4
PIN 1249/454, sin >70.8 59.0 e 6t 6.5 14.0 1.7 e 5
PIN 1249/692, sin 52.0 >70 5t 8.0 11.8 1.3 1.35 5
PIN 1249//N, dex e 62.0 e 7t 7.0 14.0 1.7 e 5
AMZ 30426/32, dex 125.0 64.8 >65.0 ~8t 7.0 13.4 1.8 1.00 5
KM 447/-379 e 82.2 >53 >6 e e 2.0 e 5
PIN 1249/723, dex 113.0 57.0 >69.0 10t 7.5 11.2 2.0 1.21 5
PIN 1249//N, sin 131.2 67.6 98.0 t9t 6.6 13.0 2.0 e 5
Sarkel (lower Don River, Russia)
SSC RAS no e 51.7 56.6 t4e 8.0 12.0 1.43 1.09 1
Port-Katon (Sea of Azov Region, Russia)
ZIN no, dex 117.0 57.0 53.0 t9t 8.0 e 1.0 0.93 3
ZIN no, sin 118.0 59.0 e t9t 8.0 e 2.0 e 4
Sredniy Egorlyk, Novo-Donskoy (Russia)
-833 e 60.2 e e4t e 11.4 1.6 e 5
Late Villafranchian
Georgievsk (Stavropol Region, Russia)
PKM 3258, dex 112.0 53.0 e t7t 7 e 1.7 e 3
SMZ no, dex 127.0 53.3 73.0 t8t 8.5 12.7 1.5 1.37
Kryzhanovka (Ukraine)
GIN no e 66.0 e >6 7.0 e 1.0 e
Middle Villafranchian
Linentsovka, Khapry (Sea of Azov Region, Russia)
AMZ 1169, sin >99 51.0 61.2 t6 7.0 14.1 1.4 1.20 3
AMZ 1446, sin 115.0 61.4 e ~7t 7.25 13.6 1.9 e 5
GIN 270/7, dex >90 61.2 66.2 t6 7.0 14.4 1.7 1.08 2
GIN 300/122, sin 120.6 64.2 e t8t 7.5 15.0 2.1 e 4
ZIN 30012, dex >91 46.8 e t7t 7.5 12.4 1.5 e 5
2853/10 108.8 57.4 e ~7t 7.5 13.3 1.6 e 4
Kuruksay (Tadzhikistan)
GIN 3848/205-73 e 76.2 65.4 6 7.0 14.2 2.0 0.86 4e5
GIN 3848, dex >96 53.6 65.0 >7 7.0 14.7 2.0 1.21 4e5
312 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318

Table 3
Tooth measurements of lower teeth dp4 of Archidiskodon meridionalis from some Villafranchian localities of southern East Europe.

dp4 Length Width Hight Dental formula (number Lamellar frequency Length of Enamel thickness Hypsodonty Wear
(mm) (mm) (mm) of plates and talons) (at 10 cm) single plate (mm) index (H/W) stage

Late VillafranchianeEarly Galerian

Sinaya Balka/Bogatyri (Taman peninsula, Russia)
PIN 1249/68, dex >115 56.2 e t9t 9.5 e e e 5
PIN 1249/92, sin >123 66.0 94.0 t7e 6.5 17.2 2.0 1.42 5
PIN 1249/99, dex >120 64.2 105.8 t7e 6.5 17.0 1.8 1.65 4
PIN 1249/168, dex >111 62.2 74.4 e7t 7.0 16.8 1.8 1.20 5
PIN 1249/no >98 55.0 e e8t 7.5 13.3 1.8 e 4
AMZ 30232/4, sin >121 60.0 90.0 t9e 7.0 15.8 2.5 e 3
AMZ 30232/14, dex >76 59.0 e t7e 7.0 13.7 2.1 e 4
AMZ 30814/1, sin 111.3 43.5 >57 t8t 9.0 12.7 e e 1
AMZ 30814/2, dex 107.35 44.3 >52 t8t 8.9 e e e 1
Tsimbal (Taman peninsula, Russia)
1802/2 P-6, sin 113.4 e >50.0 t7t e e e e 1
ZIN 25994 (28), sin 117.0 e >57 t8t 8.0 12.4 1.8 e 1
GGM 278-1, sin 151.0 54.2 84.2 t8t 6.25 15.8 1.45 1.55 4
GGM 278-1, dex 149.0 60.0 85.0 t8t 6.25 15.8 1.3 1.42 4
KM-23/P-117 97.0 52.0 e t7t 8.0 12.4 2.4 e 5
Nogaisk (Sea of Azov Region, Ukraine)
SMZ 4358/57-64, sin e 61.0 101.0 >6 6.25 15.6 1.65 1.66
Late Villafranchian
Georgievsk (Stavropol Region, Russia)
PKM 11262, sin 104.0 52.0 e t7t 7.0 e 1.7 e 3
PKM 1127, dex e 59.0 e >5 6.0 e 2.0 e 4
GIN, 78, dex 104.2 45.3 e t7t 8.0 12.4 1.64 e 4
SMZ 15399, dex 138.0 54.9 80 t8t 6.9 14.3 1.97 e 3
Saratovkaya (Psekups, Russia)
KM no, dex 113.0 62.0 e t7t 7.5 e e e 5
Middle Villafranchian
Liventsovka, Khapry (Sea of Azov Region, Russia)
ROMK X-535, sin 111.0 42.6 e t8t 6.5 15.6 e e 3
ROMK X-1185, sin 118.0 50.0 47.0 t8t 7.5 12.5 1.5 0.94 4
ROMK X-1500, sin e 58.0 e e6t 7.5 13.3 1.4 e 4
ROMK L-283, sin 132.0 57.2 75.0 t8t 7.75 14.7 1.8 1.31 4
ROMK L-910, dex >107 57.6 e ~8t 7.5 13.2 1.7 e 4
ROMK L-1366, dex e 49.0 e ~4t e e e e 5
ROMK L-1487, dex 133.0 e e t7t 8.0 14.3 e e 4e5
ROMK L-1501, sin 110.0 46.8 60.0 t8t 8.0 12.8 1.5 1.28 3
TM 147, dex >105.6 51.0 42.0 t7t e 13.2 1.6 0.82 4
TM X-5045, sin 134.0 56.0 60.0 ~7t 6.25 15.5 2.2 1.07 3
GIN 300/121, dex 141.6 61.0 >49 t8t 7.2 14.8 2.3 e 4e5

Table 4
Tooth measurements of upper teeth M1 of Archidiskodon meridionalis subspecies from some Villafranchian localities of southern East Europe.

M1 Length Width Hight Dental formula (number Lamellar frequency Length of Enamel thickness Hypsodonty Wear
(mm) (mm) (mm) of plates and talons) (at 10 cm) single plate (mm) index (H/W) stage

Late VillafranchianeEarly Galerian

Sinaya Balka/Bogatyri (Taman peninsula, Russia)
PIN 1249/006, sin 174.0 69.2 108.0 t10t 6.25 14.7 1.9 1.56 4
PIN 1249/71, sin 141.0 72.0 >89 t9t 7.25 13.6 2.2 e 4
PIN 1249/79, sin 149.6 78.0 e t10t 7.75 12.8 2.25 e 5
PIN 1249/199, sin 149.0 82.2 e ~7t 6.0 15.8 2.6 e 5
PIN 1249/237, dex 141.8 74.0 >97.0 t8e 7.0 13.1 2.6 e 4
PIN 1249/252, dex e 81.0 e t8t 6.0 15.9 2.5 e 5
PIN 1249/285, sin 145.0 70.8 >93.0 t10t 7.2 13.5 2.2 e 4
PIN 1249/699, dex 140.0 70.0 >95.0 t9t 7.0 13.7 2.3 1.36 4
PIN no, dex e 78.0 112.0 e6t 5.5 16.8 2.5 1.44 5
AMZ 30232/6, sin 145.0 72.0 >82 e8t 6.5 15.6 2.5 e 5
AMZ 30232/10, dex 139.8 71.0 e t8t 6.25 15.0 2.5 e 5
AMZ 30232/14, dex 157.0 68.2 111.0 t10t 6.0 16.2 2.3 1.63 2
AMZ 30232/15, dex 163.4 74.0 e t9t 5.75 15.4 2.5 e 5
AMZ 30232/19, dex 175.0 64.0 >93.4 t9t 6.0 15.0 e 1.46 1
AMZ 30232/22, dex e e 109.0 e3t e 15.0 e e 0
AMZ 30814/12, sin 158.0 74.8 e t8t 5.75 17.0 2.6 e 5
AMZ 30814/11, dex >157 81.2 e t8t 5.75 17.3 2.56 e 5
Tsimbal (Taman peninsula, Russia)
ZIN 25994, dex >138 87.0 e e8t 5.5 e 2.0 e 4e5
ZIN 25994 (30), sin >106 75.0 e ~6 e e 2.0 e 5
KM no, dex 143.0 85.0 e t9t 6.0 e 2.0 e 4
Locality unknown (Taman peninsula, Russia)
KM 603/P 194, sin 136.0 67.0 >66 t7t 5.25 17.0 2.8 e 4
 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318 313

Table 4 (continued )

M1 Length Width Hight Dental formula (number Lamellar frequency Length of Enamel thickness Hypsodonty Wear
(mm) (mm) (mm) of plates and talons) (at 10 cm) single plate (mm) index (H/W) stage

Nogaisk (Obitochnaya Kosa, Primorsk, Ukraine)

PRKM P-60, sin ~166 68.0 105.0 t9t 6.0 15.85 1.87 2.00 4
Port-Katon (Sea of Azov Region, Russia)
ROMK, 1042, sin 153.0 71.0 104.0 t8t 6.0 e 1.6 e
Late Villafranchian
Kotlovina (Ukraine)
ZIN 30140, dex 151.0 69.2 97.0 t8t 5.0 15.5 2.8 1.00 2
Middle Villafranchian
Linentsovka, Khapry, Kobiakova Balka (Sea of Azov Region, Russia)
ROMK X-171, dex >138 63.2 >72 ~9t 5.75 17.7 2.05 e 3e4
ROMK L-512, dex 60.0 e e 4 e e e e 4
ZIN 35733, sin >140 71.2 e ~9t 5.75 19.0 2.3 e 4
ZIN 35734, sin >142.5 72.4 e ~7t 5.75 19.2 2.1 e 5
AMZ-1171, dex 154.0 73.0 87.2 ~9t 6.5 19.0 2.3 1.19 5
AMZ-1181, sin e 70.8 e ~6e 5.25 18.4 3.0 e 5
AMZ 1324 164.0 77.0 91.0 t7t 4.7 18.2 1.8 1.18 3
AMZ-1445, dex >145 71.0 e ~7t 5.5 18.1 2.6 e 5
NMC no >157 72.6 94.8 t9t 6.25 19.5 2.5 1.31 4

Table 5
Tooth measurements of lower teeth m1 of Archidiskodon meridionalis subspecies from some Villafranchian localities of southern East Europe and adjacent territories.

m1 Length Width Hight Dental formula (number Lamellar frequency Length of Enamel thickness Hypsodonty Wear
(mm) (mm) (mm) of plates and talons) (at 10 cm) single plate (mm) index (H/W) stage

Taman Faunal complex (Late VillafranchianeEarly Galerian)

Sinaya Balka/Bogatyri (Taman peninsula, Russia)
AMZ 29899/2, dex 169.0 75.0 >44 t10t 6.5 15.3 2.0 e 4
AMZ 29899/7, dex 160.0 70.8 >92.0 t10t 6.0 14.0 2.2 1.30 4e5
AMZ 30426/37, dex 152.2 60.2 e t9t 6.5 15.3 2.0 e 5
AMZ 30232/2, dex 144.0 70.0 e t8t 6.5 15.5 2.1 e 5
AMZ 30232/5, sin 150.2 72.2 107.0 t9t 7.0 15.6 2.5 1.48 5
AMZ 30232/16 >139 70.2 e t9t 6.25 14.8 2.3 e 5
AMZ 30814/5, dex 149.0 72.0 >90.7 t9t 7.125 17.0 2.0 1,2e592 4e5
AMZ 30814/8, sin 147 61.9 e t8t 7.125 15.0 2.5 e 3
PIN 1249/65, sin e 67.0 e e4t 5.5 18.2 2.3 e 5
PIN 1249/144, sin 148.0 68.0 102.2 t8t 5.75 16.6 2.1 1.50 5
PIN 1249/169, sin >131 79.0 >76.0 ~8t 6.0 17.1 2.6 0.96 5
PIN 1249/210, sin >127 72.4 e e6t 5.0 19.7 2.6 e 5
PIN 1249/253, dex 169.0 75.0 e t9t 5.0 17.6 2.3 e 5
PIN 1249/677, dex >119 69.0 >82.0 e7t 5.25 19.5 2.5 1.19 3
PIN 1249/678, dex >120 66.8 >88.4 t8e 6.75 16.0 1.7 1.32 5
PIN 1249/682, sin >126 72.0 >77.0 e7t 5.5 17.9 2.3 1.07 5
PIN 1249/681, sin >131 60.0 >94.6 t8e 6.25 17.2 2.1 1.58 4
PIN 1249/701, dex e 67.6 e ~4t 5.5 19.5 2.0 e 5
PIN 1249/730/733, sin 158.0 67.0 104.0 t8t 6.0 17.3 1.9 1.55 4
GGM 0847-11, dex 170.5 74.0 e t8t 5.0 19.5 2.13 e 5
Tsimbal (Taman peninsula, Russia)
GGM 278-1, dex 149.0 60.0 85.0 t8t 6.25 15.82 1.3 1.42 4
GGM 278-2, sin 151.0 54.2 84.2 t8t 6.25 15.85 1.45 1.55 4
Port-Katon (Sea of Azov Region, Russia)
PIN 2057, sin, dex 162.0; 163.0 72.0; 73.0 88.0 e8t 5.5 e 2.2 1.31 4
Samarskoe (Sea of Azov Region, Russia)
ROMK 1121, dex 143.0 63.2 >64 t7t 6.25 15.7 2.2 e 5
ROMK 1248, sin 151.0 62.0 82 t8t 6.5 e 2.0 1.32 2
Khapry faunal complex (Middle Villafranchian)
Linentsovka, Khapry, Mokriy Chaltir' (Sea of Azov Region, Russia)
GIN 270/6, dex >135 60.8 e ~7t 6.25 16.3 2.1 e 5
GIN 300/17, dex 142.0 66.8 e t8t 5.5 17.2 2.3 e 5
ROMK L-86, dex >141 59.0 e ~8t 6.0 17.6 2.0 e 4
ROMK L-190, sin e 62.0 e e7t 6.25 15.5 2.1 e 4
ROMK L-512, sin e 66.0 e t3e 5.5 e 1.9 e 4
ROMK L-910, dex 140.2 56.0 77.0 t7t 6.5 15.5 1.9 1.37 0
ROMK L-979, sin e 55.8 73.0 t5e 6.5 e 1.9 1.31 1
ROMK L-1366 149.0 59.0 87.8 t8t 6.5 17.9 e 1.49 3
ROMK L-1467 e 64.0 e t5e 6.0 e 1.9 e 4
ROMK X-1353, dex 145.8 59.8 87.0 t8t 5.5 18.2 2.0 e 4
ROMK MCH-1353, dex >140 59.0 >70.0 ~8t 5.5 17.5 2.0 1.19 4
AMZ, 1453, sin 150.0 61.0 e t8t 6.25 15.7 2.2 e 4
AMZ MCH no, sin 138.1 62.6 70.7 t8t 6.0 17.1 2.6 1.13 3
314 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318

Table 6
Tooth measurements of DP4/dP4 e M1/m1 from Early Pleistocene elephants from Eastern Europe: Sinaya Balka/Bogatyri site (Late VillafranchianeEarly Biharian), and Liv-
entsovka, Khapry localities (Middle Villafranchian). Data in brackets are mean. s e standard deviation.

Measurements Archidiskodon m. tamanensis, Sinaya Balka Archidiskodon m. gromovi, Liventsovka, Khapry

dP4/4 M1/1 dP4/4 M1/1

n min-(med)-max s n min-(med)-max s n min-(med)-max s n min-(med)-max s

Length, mm 6 113:0126:2134:0 7:3 13 139:8152:20175:0 12:4 3 108:8114:8120:6 5:9 2 154:0;160:0 e
4:8
3 107:35116:8131:9 13:2 14 142:3154:0170:5 9:6 6 110:0123:0134:0 12:5 6 138:0144:2150:0
Width, mm 15 50:460:4774:2 6:2 18 64:074:5086:1 5:8 6 46:857:064:2 6:8 9 60:068:9173:0 4:7
11 43:556:1166:0 7:1 24 60:069:179:0 5:0 10 42:648:6758:0 5:7 13 56:060:9166:8 3:4
Height, mm 3 88:497:8107:0 9:3 5 93:4106:68112:0 7:6 2 61:2; 66:2 e 3 85:089: 094:8 5:1
2 94:0; 105:8 e 4 102:0104:4107:0 2:0 2 60:0; 60:2 e 5 71:079:187:8 7:9
Number of plates (including talons) 5 1010:411 0:55 12 1011:012 0:85 3 99:3310 0:6 3 1010:6711 0:6
4 1010:511 0:6 14 1010:712 0:7 4 99:7510 0:5 6 1010:010 0:0
Number of plates (excluding talon) 5 88:49 0:55 12 89:010 0:85 3 77:338 0:6 3 88:679 0:6
4 88:59 0:6 14 88:710 0:7 4 77:758 0:5 6 88:08 0:0
Lamellar frequency 15 6:57:178:0 0:5 17 5:46:317:75 0:7 6 7:07:297:5 0:25 8 5:255:786:5 0:4
9 6:57:859:5 1:2 24 5:06:27:1 0:7 9 6:257:448:0 0:6 13 5:56:026:5 0:4
Length of single plate, mm 15 11:213:0614:5 1:0 17 12:815:0817:3 1:4 6 12:413:815:0 0:9 8 17:718:8119:6 0:7
7 11:8514:6517:2 2:3 24 13:916:619:7 1:8 11 12:513:9215:6 1:1 10 16:016:8518:2 1:0
Enamel thickness, mm 15 1:31:762:2 0:3 16 1:92:392:6 0:2 6 1:41:692:1 0:3 7 2:052:392:85 0:3
8 1:81:882:1 0:1 24 1:72:22:6 0:3 9 1:41:72:2 0:3 12 1:82:052:6 0:2
Hypsodonty index, HI (H/W) 2 1:57; 1:62 e 4 1:441:521:63 0:1 2 1:08; 1:2 e 3 1:191:261:31 0:1
2 1:42; 1:65 e 3 1:51:511:6 0:0 2 1:07; 1:28 e 4 1:11:331:49 0:15
Crown proportion index (W/L) 6 45:350:4855:4 3:4 12 36:5747:6055:17 5:45 2 52:76; 53:23 e
1:8
2 41:25; 47:4 e
1
3 39:140:9942:61 1:8 10 39:645:649:1 3:1 5 38:441:6843:3 6 40:042:2747:0

the previous teeth generation. Teeth of Taman elephant are

The digital information was processed with the help of com-
puter programs MS Excel and Statistica 8.0. We used the Shapir-
oeWilk test (SeW) for the normality of parameter distribution
verication.
3. Description and comparison
The preservation degree of the material from Sinyaya Balka/
Bogatyri is more or less homogeneous (Figs. 5 and 6). The dental
cement is usually brown (50%) or cream (41.7%), rarely creamy
brown (8.3%), sometimes covered with manganese oxide dendrites.
Tooth enamel is predominantly gray (40%) and grayebrown
(33.3%), blueegray (13.3%), creamy blue (6.7%) and light cream
(6.7%). Dentin is usually brown (46.15%), white (38.5%), or more
rarely creamy brown (15.35%).
We have a sample of teeth DP4/dp4 e M1/m1 from site Sinyaya
Balka/Bogatyri, which is sufcient for analysis. Therefore, we made
a detailed description of parameters of these teeth generations
only. It allows identication of their characteristics and variability
of features.
Teeth M1/m1 have a longer crown than the DP4/dp4 by average
and by minimal and maximal values. Length of DP4/dp4 varies
within the limits of 107.4e134 mm, and M1/m1 e 140e175 mm.
However, the determination of early generation teeth order should
be done in combination with other characteristics, as the values of
crown length overlap signicantly. The numerical measures of
teeth length of A. m. tamanensis and A. m. gromovi overlap and differ
by the average values only. This parameter insignicantly varies
from those on teeth of M. trogontherii, and on average is larger than
for European M. primigenius (Garutt and Foronova, 1976; Dubrovo,
2001; Maschenko, 2002).
Mean values of upper teeth's crown width exceed those of the
lower ones, both on DP4/dp4, and on M1/m1 (Table 6). On average,
the values of this parameter for elephants from Sinyaya Balka
exceed those for A. m. gromovi from Liventsovka and Khapry.
However, they coincide with the dimensional characteristics of the
teeth of M. trogontherii from Sssenborn, and on average are larger
than for M. primigenius.
Despite the signicant wear in the greater part of the studied
specimens, we can note some patterns in crown height. The crown
height of upper teeth is slightly larger than those of lower ones for
elephants from Sinyaya Balka/Bogatyri and from Liventsovka and
Khapry (Table 6). On M1/m1, the height is slightly greater than for
noticeably higher than those of Gromov's elephant.
The number of enamel plates on DP4/dp4 and on M1/m1 is the
same for the upper and lower teeth. Such regularity was marked for
M2/m2 and M3/m3 of the Taman elephant (Dubrovo, 1960). The
number of plates is 10 or 11 with talons on teeth DP4/dp4 (dental
formula is t8t (n 5) and t9t (n 3)). The majority of M1/m1 has
the same number of plates as those for the previous generation, 10
(t8t (n 10)) and 11 (t9t (n 10)). Six specimens have 12 plates
(t10t).
On the A. m. gromovi sample from Khapry Faunal complex lo-
calities, this characteristic is smaller by one plate (Table 6). Four
ndings of DP4/dp4 have dental formula of t7t and the same
number of ndings have a formula of t8t. The majority of M1/m1
consists of 10 plates with talons (dental formula t8t (n 10)) and
only two specimens include 11 plates (t9t). Teeth of elephants from
the Sinyaya Balka/Bogatyri are smaller than those of the trogon-
therii elephants from various Middle Pleistocene localities of
Europe by 1e2 plates (Garutt and Foronova, 1976; Dubrovo, 2001).
Taking into consideration that according to majority of researchers
the number of plates that make up the crown is one of the most
diagnostic feature of elephants, this indicator reects an adequate
picture of gradual evolutionary change not only at the last teeth
generation, but in earlier generations as well.
On teeth DP4/dp4 from Sinyaya Balka/Bogatyri, the values of
lamellar frequency at 10 cm, one of the most diagnostic features,
range from 6.5 to 9.5 and from 4.5 to 7.1 on the M1/m1. On average,
these measures are higher than those on teeth of A. m. gromovi from
Liventsovka and Khapry (Table 6), but less than the Middle Pleis-
tocene M. trogontherii, and signicantly less than those of Late
Pleistocene M. primigenius from European localities. The lamellar
frequency is larger on lower dp4-m1 and has a greater range of
variability. This is due to a stronger divergence of plates to the
crowns' base on lower teeth due to increased length in ventral di-
rection and bending of the tooth (Dubrovo, 1960).
Length of enamel plates decreases from the anterior to the
posterior part of the crown and depends on the angle and degree of
tooth wear. It varies quite widely from 11.2 to 17.2 mm on DP4/dp4
and from 14.0 to 19.7 mm on M1/m1. These features indicate that
the plates of elephant teeth from Sinyaya Balka/Bogatyri, on the
average, are thinner than those from Liventsovka and Khapry, but
thicker than those of trogontherii elephants.
The enamel thickness of the dental plates ranges from 1.3 to
2.2 mm on DP4/dp4 and from 1.7 to 3.0 mm on M1/m1 (Table 6).
 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318
The enamel is weakly plicated on weakly eroded teeth and middle
folded on medium- and heavily worn ones. The mean values of
enamel thickness on teeth from Sinyaya Balka/Bogatyri coincide
with those from Liventsovka and Khapry, but slightly higher than
the values of M. trogontherii from Sssenborn and Tiraspol.
According to Dubrovo (1960) one of the diagnostic feature of last
generation teeth M2 and M3 of A. m. tamanensis is the incipient
wear gures on the occlusal surface of trinomial plates at the level
of 10e20% of their height. According to her description, this gure
is usually of antiquoid type (  ) or similar to it. We have seen 39
DP4/dp4  M1/m1 from Sinyaya Balka/Bogatyri in order of wear
stage of the crown e from weak wear degree (0 or 1 stage) to
signicant (5 and 6 wear stages). In 84.6% of cases, the sample in-
cludes teeth with antiquoid wear, 10% e similar to those of
M. trogontherii from Tiraspol (  ), and 5.4%, a meridionaloid
gure (  ). Compared with the sample of 20 DP4/dp4 and M1/
m1 of the Gromov's elephant from Liventsovka and Khapry, only
30% of specimens have the antiquoid gure, the meridionaloid
gure is prevalent (65%), and only 5% have an intermediate type for
M. trogontherii. Often, there are median sinuses of enamel at the
anterior and posterior sides of enamel plates on the teeth of the 4th
and 5th wear stages.
Dubrovo (1960) noted that taxonomic position of elephants
cannot be determined on a basis of incipient wear gures of plates.
It is necessary to take into account variability in this feature at
different taxa. Two subspecies of southern elephant A. m. tam-
anensis and A. m. gromovi lived in the same area over more than one
million years. The noted differences of plate wear gures may
reect changes in these animals' diet as a result of gradually
changing living conditions. However, the wear gure of three
anterior plates on the teeth from Sinyaya Balka/Bogatyri is mainly
of mixed type (  ), and it is usually antiquoid, beginning with
the fourth one.
For Sinyaya Balka/Bogatyri teeth, the hypsodonty index, on
average, is slightly higher than the same for Khapry elephants
(Table 6). Its value ranges from 1.0 to 1.62 on DP4, from 1.2 to 1.65
on dp4, from 1.36 to 1.63 on M1, and from 1.0 to 1.6 on m1.
The range of the standard deviation of the parameters (Table 6)
shows that dimensions of the teeth (length, width and height of the
crown) are signicantly changeable due to the wear degree. Stan-
dard deviations of the parameters of lamellar frequency and
number of plates, and enamel thickness, usually do not exceed 1.0.
However, standard deviation of the length of single plates is greater
than 1.0, which makes it possible to divide the estimated taxa of the
sample by this parameter.
4. Discussion and conclusion
The examined series of deciduous premolars and rst molars
from Sinyaya Balka/Bogatyri show that there is a poor ability to
detect different forms of meridionaloid elephants on the basis of
data of milk teeth DP2/dp2 e DP3/dp3, due to insufcient sample
size. The comparison of DP4/dp4 parameters distinguishes
different morphotypes in one collection with a greater validity and
recognition of some differences in the characteristics of teeth of the
southern elephants from Sinyaya Balka/Bogatyri and Liventsovka
based on the average and optimal characteristics. Features of M1/
m1 reveal more differences between samples from different age
localities. Despite the lower diagnosticity of teeth DP4/dp4 e M1/
m1 compared with M3/m3, our studies have demonstrated the
feasibility of their use for taxonomic denitions. The prevalence of
more advanced features on the teeth of A. m. tamanensis (Late
VillafranchianeEarly Biharian) compared to A. m. gromovi (Middle
Villafranchian) was noted. The most revealing ones were the
graphics on the upper DP4 and M1 (Fig. 5). They make it possible to
315
distinguish the teeth of southern elephants from localities of early
and late Early Pleistocene of Eastern Europe quite clearly. The
graphics on the lower teeth were less informative. Apparently, this
is due to the fact that the analyzed parameters of lamellar fre-
quency and length of a single plate considerably change on lower
teeth depending on the wear stage, more than on the upper ones.
The average length of single plate is a very diagnostic characteristic
of early generation teeth.
Teeth parameters of Late Villafranchian A. m. meridionalis fall
between A. m. tamanensis and A. m. gromovi. The described teeth
generations reliably distinguish the meridionaloid elephants from
the later representatives of mamontoid lineage elephants, for
example, from Middle Pleistocene Mammuthus trogontherii and
M. chosaricus, and certainly from the Late Pleistocene
M. primigenius.
The available teeth series conrms the results of the sample's
heterogeneity from localities of Taman fauna from south Eastern
Europe, previously made on the basis of studying the teeth of last
changes (Lister and Sher, 2001; Lister et al., 2005; Baigusheva and
Titov, 2012). This is conrmed by the absence of normal distribu-
tion of features of DP4/dp4eM1/m1 in the collection from the
Sinyaya Balka/Bogatyri (Fig. 7). We did not observe a clear bimodal
distribution of main diagnostic characteristics in the studied
collection, which could be supposed in the case of the presence of
two taxa. According to the ShapiroeWilk test (SeW) criterion
(Fig. 7), with values W < 1 (W values vary within 0.68e0.98 for
different parameters), the hypothesis of normal distribution of the
studied variables can be rejected. However, the levels of statistical
signicance of the samples are mostly p > 0.05. This suggests that
the distribution is not very different from normal. By this criterion,
the most signicant differences in the sample have parameters of
lamellar frequency (LF), number of plates (NP), and the length of
single plate (LSP) (in case of more than 10 specimens). This assumes
an occurrence of admixture of separate specimens of other taxa in
the bulk of Taman elephant nds, which are more numerous.
However, we could not reliably determine the taxonomic afliation
of the teeth of the second form, considering the rather insufcient
diagnosticity of the investigated teeth generations, transgressive
overlapping of features, and probable similarity of Early Pleistocene
elephants' dental characteristics. Some teeth (e.g., specimens M1
No. PIN 1249/006 and AMZ 30232/19) show similarity with the
holotype of Phanogoroloxodon mammonthoides (Fig. 8). These teeth
have relatively thin enamel, long and narrow tooth crowns, but
they are not very different from the main group of Taman elephants
in the number and frequency of plates (and therefore the length of
single plates). In addition, the tooth M1 of holotype of phanagorian
elephant has a small pathology of 2 and 3 enamel plates on the
lingual side of the crown (Fig. 3). This complicates the comparison.
It is conditionally possible to allocate some teeth as DP4/dp4,
and M1/m1 with a slightly higher lamellar frequency and length of
single plate among other ndings from Sinyaya Balka/Bogatyri.
These specimens are close to the lower limit of variability of the
same teeth of M. trogontherii by their characteristics, although they
do not coincide completely. When analyzing a nature of variability
at the wearing process, teeth with greater wear (the 5th stage),
usually have thicker enamel, smaller lamellar frequency and
greater length of single plates (Fig. 8). Perhaps this is one of the
factors that increase a degree of parameter dispersion and explains
the absence of clear normal distribution of characteristics.
The possibility of re-deposition or condensation of remains of
animals of different ages in the Sinyaya Balka/Bogatyri site is un-
likely. This locality has a complex geological situation and struc-
ture, but it is considered to be coeval, although three horizons of
somewhat differing lithology and genesis are separated in its
structure. The two upper ones contain the remains of large
316 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318

Fig. 7. The frequency distribution of parameters of teeth DP4/dp4 and M1/m1 of elephants from locality Sinyaya Balka/Bogatyri (Taman peninsula, Russia): lamellar frequency (LF),
length of single plate (LSP), enamel thickness (ET), number of plates without talons (NP), and a ratio of width to length (W/L).

Fig. 8. Ratio of enamel thickness and length of single plate of upper teeth M1 and DP4 of different age subspecies of Archidiskodon meridionalis from Eastern Europe and adjacent
territories.
 V.S. Baigusheva et al. / Quaternary International 420 (2016) 306e318
mammals. However, the greatest amount of bone (90%) occurs
from the latest and most notable of these layers. Paleontological,
lithological, and archaeological evidence suggests that the for-
mation of the locality was not instantaneous, but within one age
stage (Shchelinsky et al., 2010). According to the level of evolu-
tionary development, the remains of large mammals, as well the
species compositions of the fauna, are not different for these two
horizons.
At this stage of research, based on the studying of the sample of
early generations' teeth of Taman elephant from Sinyaya Balka/
Bogatyri, we assume that the bulk of the collection is represented
by remains of one elephant taxon, A. m. tamanensis. Taking into
consideration that meridionaloid elephants in the Early Pleistocene
of the Northern Caucasus coexisted with phanagorian elephant Ph.
mammonthoides, we do not exclude the possibility that separate
nds belong to this form of elephant, which has similar dental
characteristics (Fig. 8).
However, the examined dental material does not allow us to
reliably diagnose these samples with some different parameters.
Further study of all teeth generations, bones of the postcranial
skeleton from the locality, as well as the possible discovery of skulls
on which these species are easily distinguished, will clarify the
issue of the taxonomic composition of Proboscideans from Sinyaya
Balka/Bogatyri.
Acknowledgements
The authors are grateful to E.N. Maschenko, A.K. Aghadjanian
(Paleontological Institute, Russian Academy of Sciences, Moscow),
I.L. Soroka, I.A. Starodubtseva (State Geological Museum. VI Ver-
nadskiy, Moscow), A.S. Tesakov, P.A. Nikolsky (Geological Institute,
Russian Academy of Sciences, Moscow), A.K. Shvyreva (Stavropol
State Museum-Reserve, Stavropol) for the opportunity to study the
collections. The authors are particularly grateful to employees of
the Azov historical-archaeological and paleontological museum-
reserve (Azov) G.I. Timonina and S.V. Semionova for their help in
the excavation and processing of nds from locality Sinyaya Balka/
Bogatyri. We are grateful to V.E. Shchelinsky and S.A. Kulakov
(Institute of History of Material Culture, St. Petersburg) for their
assistance in the investigation of the site. Special thanks to Dr. A.
Chikin (IAS SSC RAS, Rostov-on-Don) for consultations on statistical
analysis of the material. This work was supported by Russian fund
of basic researches No. 15-04-02079-a and 15-05-03958-a.
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