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Palaeontologia Electronica
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Lobachev, Y.V., Shpansky, А.V., Bondarev, A.A., Lobachev, A.Y., Vasiliev, S.K., Klementev, A.M., Grebnev, I.E., and Silaev, V.I. 2021.
New findings of Stephanorhinus kirchbergensis in Siberia. Palaeontologia Electronica, 24(1):a14. https://doi.org/10.26879/734
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New findings of Stephanorhinus kirchbergensis in Siberia
Y.V. Lobachev, А.V. Shpansky, A.A. Bondarev, A.Y. Lobachev, S.K. Vasiliev,
A.M. Klementev, I.E. Grebnev, and V.I. Silaev
ABSTRACT
New findings of Stephanorhinus kirchbergensis (Jäger, 1839) remains, obtained
from the Asian part of Russia, are described. The material includes 39 specimens from
13 localities in West Siberia and East Siberia. It considerably expands the geographic
distribution of this species of rhinoceros. A series of 11 mandibles from Siberia, includ-
ing one juvenile individual with deciduous teeth, is described for the first time. We also
present a large set of data on well-preserved postcranial remains. The morphology and
sizes of mandibles, teeth, and postcranial remains of adult individuals of S. kirchber-
gensis from Siberia are similar to individuals of this species described from European
localities. A series of upper teeth was subjected to mesowear analysis to assess the
diet of S. kirchbergensis from West Siberia. The chemical composition (including stable
isotopes) of the Siberian Stephanorhinus teeth is analyzed for the first time. Compari-
sons of Siberian S. kirchbergensis with European S. kirchbergensis and West Siberian
Coelodonta antiquitatis broaden our understanding of the ecology, variability, and evo-
lution of S. kirchbergensis under climatic changes in continental settings from the Mid-
dle to the Late Pleistocene. Despite small samples, we can suppose that S.
kirchbergensis was widely distributed in Siberia.
Y.V. Lobachev. PAO Novosibirsk Institute of Software Systems, Novosibirsk, Russia.
yvlobachev@gmail.com
А.V. Shpansky. Tomsk State University, Russia. shpansky@ggf.tsu.ru
A.A. Bondarev. Omsk Regional Branch of the Russian Geographical Society, Omsk 644007, Russia.
gilgamesh-lugal@mail.ru
A.Y. Lobachev. PAO Sberbank, Novosibirsk, Russia. inobges@gmail.com
S.K. Vasiliev. Institute of Archaeology and Ethnography of SB RAS, Russia. Svasiliev@archaeology.nsc.ru
A.M. Klementev. Institute of the Earth’s Crust SB RAS, Russia. klem-al@bk.ru
I.E. Grebnev. Paleopark Altai Republic, Russia. mnh66@mail.ru
V.I. Silaev. Institute of Geology of Komi SC UB RAS, Syktyvkar, Russia, silaev@geo.komisc.ru
Keywords: Stephanorhinus kirchbergensis; bone morphology; Siberia; Middle Pleistocene; mesowear
analysis
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
2
Submission: 21 October 2016. Acceptance: 15 May 2021.
INTRODUCTION
Remains of Stephanorhinus kirchbergensis
(Jäger, 1839) are relatively rare. There are about
80 localities known, most of them being in the
European part of the geographic distribution. Only
a few remains of S. kirchbergensis have been
described from the Yakutia (Sakha) Republic and
the Irkutsk, Kemerovo, and Tomsk Provinces, Rus-
sia (Billia, 2007, 2008b, 2011a, 2011b). The only
skull from Siberia lacks georeferenced data but
presumably came from the Irkutsk Province (Cher-
sky, 1874; Billia, 2008a). Dubrovo (1957) described
S. kirchbergensis teeth (previously Rhinoceros
mercki) and Mammuthus trogontherii remains (pre-
viously Parelephas wüsti) from the Vilyuy River
(Yakutia). Isolated teeth of S. kirchbergensis were
reported in Mohovsk Quarry and the Inya River of
the Kemerovo Province (Billia, 2007). Most S.
kirchbergensis remains from Russia, more than 30
teeth and bones, were found in Krasniy Yar (Tomsk
Province) (Alekseeva, 1980; Shpansky and Billia,
2012; Shpansky, 2016). All of the previously
described material from Siberia was assigned to
the Middle Pleistocene (Volkova and Babushkin,
2000).
In the last few years, a large series of new
findings of Stephanorhinus kirchbergensis was
obtained from southern West Siberia, including the
Altai and Krasnoyarsk Territories and the Novosi-
birsk, Omsk, Tomsk, and Irkutsk Provinces. The
new material, described here for the first time,
came from 13 localities in the southern part of West
Siberia and East Siberia (Figure 1). The material
consists of 39 samples, including a series of 11
mandibles. One mandible belongs to a juvenile
with deciduous teeth. Most of the samples did not
have a stratigraphic provenance as they were col-
lected on riverbanks.
MATERIAL
Altai Territory
A collection of 5,806 bones, which were
attributed to 24 species of mammals of the Middle
and Late Pleistocene, was gathered on river
beaches and shallows of the Chumysh River
between the villages Martynovo and Kytmanovo
since 2010. The most numerous remains were
found near Novoduplenka village (53º27' N, 85º41'
E) (Figure 1). The findings near Novoduplenka
include 19 very well-preserved specimens
assigned to Stephanorhinus kirchbergensis: three
mandibular corpora (including one belonging to a
juvenile individual with deciduous teeth), one verti-
cal mandibular ramus, eight upper teeth, and
seven postcranial bones (Table 1) (Lobachev et al.,
2014; Vasiliev et al., 2014).
In the study area, the Chumysh River
exposed the deposits of low first and second fluvial
terraces, which were from 12 to 20 m high. Radio-
carbon dating of wood remains from the lowest part
of five sections in the valley of the Chumysh River
(Kytmanovo, Staroglushinka, Sorokino, Pogorelka,
Shadrintsevo) indicate a Karginian age within 38–
24 14C ka BP (Rusanov and Orlova, 2013). Thir-
teen bones collected on the beaches between Mar-
tynovo and Kytmanovo were sampled for
radiocarbon dating (no bones of Stephanorhinus
kirchbergensis were sampled for radiocarbon dat-
ing). Four dates presented in a report (Lobachev et
al., 2012) and nine unpublished dates indicate the
age of the sampled fossils ranges from 44.5 to 22.3
14C ka BP. Taphonomic observations of the speci-
mens by one of us (AVS) indicate that all mandi-
bles (both juvenile and adult) were slightly
rounded, which could reflect redeposition from
more ancient deposits of the Middle to Upper Pleis-
tocene. On this part of the Chumysh River, locali-
ties with deposits that suggest an older age are
known, including clam shells (SB RAS-425) from
base layer 6 in the settlement of Kytmanov (Rusa-
nov and Orlova, 2013).
Two mandibles and two postcranial bones
were found on the sandy shoals in the vicinity of
Biysk (the exact location is unknown) (Table 1). It is
assumed that these remains were washed away
from the base layer of high terraces of the Biya
River (Rusanov and Orlova, 2013). These natural
outcrops are known near the village Staraya
Azhinka (the height of the terrace is 64 m), 60 km
to the east of Biysk, where alluvial deposits of the
Middle–Upper Pleistocene are exposed. Some
bones collected from the middle height of the ter-
race did not contain collagen (SB RAS-4003).
Another outcrop, a 50 m high terrace, is on the
right bank of the Biya River, upstream from
Stanitsa Bahtemiskaya village, 30 km east of
Biysk.
PALAEO-ELECTRONICA.ORG
3
Toms k Provin ce
New remains from the Tomsk Province were
found in the well-known locality Krasniy Yar (57º07'
N, 84º30' E) (Alekseeva, 1980; Shpansky and Bil-
lia, 2012; Shpansky, 2016; Shpansky et al., 2016),
as well as from Asino and Kindal (Figure 1). New
findings comprise a mandibular corpus, an upper
tooth, and four postcranial bones (Table 1). The
geological age is assumed to be Tobolian (MIS 11-
9). The AMS radiocarbon analysis (UBA-21200
and UBA-21201) carried out on an astragalus of
Stephanorhinus kirchbergensis (PM TSU 5/740)
showed the absence of collagen in the bone
(Shpansky et al., 2016).
FIGURE 1. New findings of Stephanorhinus kirchbergensis remains in Western and Eastern Siberia. Red - new local-
ities and finds, yellow - previously described localities. 1, Kytmanovo. 2, Biysk. 3, Asino. 4, Kindal. 5, Krasniy Yar,
Tomsk Province. 6, Berdsk. 7, Krasniy Yar, Novosibirsk Province. 8, Bibiha. 9, Taradanovo. 10, Utuskun. 11, Omsk.
12, Kachulka. 13, Irkutsk Province.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
4
A tooth (PM TSU 1/396) was found at a 8 m-
depth in Tobolian deposits in a sand quarry east of
Asino on the left bank of the Chulym River
(57º04'N, 86º10'E).
The most remarkable specimen is a mandible
(KB MAN K-397) found at 14 m below water level
at the mouth of the Kindal oxbow, left bank of the
Ob River, downstream from the Kargasok settle-
ment in July 2011 by the local resident A.V. Bary-
TABLE 1. Material included in this study.
Locality Skeletal element Collection Name
1 Kytmanovo, Chumysh River, Altay Territory right М2 NSMLL-1
right Р4 NSMLL-2
right М3 NSMLL-3
left М3 NSMLL-4
left М2 NSMLL-5
left Р4 NSMLL-6
left Р4 NSMLL-7
left Р4 NSMLL-8
left metatarsal II NSMLL-101
left metacarpal II NSMLL-102
left metatarsal IV NSMLL-105
astragalus NSMLL-107
right radius IAE CHU-108
left ulna NSMLL-109
conjoined left tibia and fibula NSMLL-110
right body of mandible NSMLL-10
left body of mandible, juvenile NSMLL-12
right body of mandible NSMLL-26
right ramus of mandible NSMLL-27
2 Biysk vicinity, Altay Territory right body of mandible GR PC-1164
mandible GR PC-1165
left metacarpal II GR PC-203
right metacarpal II GR PC-214
3 Asino, Chulym River, Tomsk Province left M1 PM TSU 1/396
4 Kindal, Ob River, Tomsk Province left body of mandible KB MAN K-397
5 Krasniy Yar, Ob River, Tomsk Province metacarpal III PM TSU 5/5197
Navicular PM TSU 5/2538
Navicular PM TSU 5/3063
6 Berdsk vicinity Novosibirsk Province third left upper molar M3 IAE BB-1
7 Krasniy Yar, Ob River, Novosibirsk Province left body of mandible NSMLL 22090
left body of mandible IAE KY-4323
8 Bibiha, Ob River, Novosibirsk Province right М2 NSMLL 21052
9 Taradanovo, Ob River, Novosibirsk Province left metacarpal IV IAE TRD-17
left metacarpal IV IAE TRD-18
left metatarsal IV IAE TRD-2
10 Utuskun, Irtysh River, Omsk Province right body of mandible Sk_ui1
11 Omsk, Irtysh River, Omsk Province left р4 Sk_oms1
12 Kachulka, Krasnoyarsk Territory left р3 PM TSU № 1/395
13 Irkutsk Province mandible IRM 2436
PALAEO-ELECTRONICA.ORG
5
shev. The locality is 400 km north of the city of
Tomsk (Kargasok area of the Tomsk Province).
The Kindal location of Stephanorhinus kirchber-
gensis is 59º08' N, 80º35'E, which is closer in lati-
tude to the finding on the Vilyuy River in Yakutia
(63º40' N) (Dubrovo, 1957).
Novosibirsk Province
Mandibular corpora of two individuals of
Stephanorhinus kirchbergensis (NSMLL 22090
and IAE KY-4323) were discovered in the Krasniy
Yar locality (55º13' N, 82º52' E) on the left bank of
the Ob River, 17 km north of the city of Novosibirsk
(Figure 1). These remains were found among
3,336 bones of other species of large mammals
that had inhabited the southeastern area of West
Siberia in the Middle–Late Pleistocene. Samples
were collected on the sandbank. Some of them
wear marks of redeposition. Analysis of spores and
pollen, seeds, ostracods, and mollusks revealed
warm interglacial conditions at the time of layer
deposition. Kazantsevo Horizon (MIS 5) is the
most probable origin for the alluvial deposits of the
channel of layer 6 (Martynov et al., 1977; Volkov
and Arhipov, 1978; Panychev, 1979). A high pro-
portion of the remains from this locality may have
been redeposited, which suggests a considerable
range in age. Thus, Middle Pleistocene age for S.
kirchbergensis remains should not be excluded.
An upper molar (NSMLL 21052) was found
near Bibiha village (55º19' N, 82º52' E) on the right
bank of the Ob River, 26 km north of Novosibirsk
(Table 1). This locality is represented by cross-bed-
ded sands of the outcrop of the third fluvial terrace
sloping below the water surface, similar to those at
Krasniy Yar (Novosibirsk Province), layer 6.
An upper left molar (IAE BB-1) was found
during dredger activities within the city limits of
Berdsk (Table 1). The exact location is unknown.
Three rhinoceros metapodials were discov-
ered among the numerous large mammalian
remains from Taradanovo (53º48'N, 81º49'E) on
the Ob River (Suzun area of the Novosibirsk Prov-
ince) (Figure 1, Table 1). All megafaunal remains
are redeposited. The main bone-bearing horizon of
the Taradanovsk highbank is located several
meters below the lowest shore line. The section of
highbank near Taradanovo has a similar structure
to that at Krasniy Yar near Novosibirsk. Eighteen
radiocarbon datings were obtained from the bones
collected from the beach: 13 dates indicate an age
older than 45–40 14C ka BP, which is close to the
limits of the method, and five dates indicate an age
of 36–26 14C ka BP (Vasiliev and Orlova, 2006).
The diversity of fauna includes the species of the
beginning of Late Pleistocene, which allows esti-
mating its geological age. One of us (SKV) thus
supposes that the main bone-bearing layer in Tara-
danovo was formed during the Kazantsevian (MIS
5).
Omsk Province
A mandibular corpus (Sk_ui1) was found by
local historian N. B. Peristov in the Irtysh River
near Krasnoyarka and Utuskun villages of the Ust-
Ishymsk district, Omsk Province (57º44'N, 71º17'E)
(Figure 1). The Ust-Ishymsk area includes a group
of early–Late Pleistocene mammal-bearing locali-
ties, typical for the area of latitudinal flow of the
Irtysh River. Terrace deposits of Karginian and Sar-
tanian ages are located in the vicinity of the site
(Krivonogov, 1988). Up and down the stream of the
Irtysh River, the outcrops of lower and Middle
Pleistocene are exposed on the right bank. Lower
Pleistocene sediments are observed in the out-
crops of the upland “Tobolsk continent”: the Koltyr-
minsk, Narimanovsk, Nikolsk, Romanovsk, and
Sakanairsk cliffs.
A lower fourth premolar (Sk_oms1) was found
by local historian A. L. Dorogov in Omsk on the
bank of the Irtysh River. A lithological description of
natural outcrops of the valley became impossible
due to anthropogenic activities. Mammal bones of
various ages from the Pliocene to the present
occur within the city on the sandbanks and in artifi-
cial outcrops.
Krasnoyarsk Territory
A lower third premolar (PM TSU 1/395) was
found in Kachulka village (53º47'N, 92º54'E),
Karatuzsk district, Krasnoyarsk Territory, 100 km
east of Minusinsk at the confluence of the Amyl
and Kazyr rivers (Figure 1, Table 1). The tooth was
buried in fine-grained sands at a depth of about 3
m. Its preservation suggests redeposition from
older deposits. This finding in the south of the
Krasnoyarsk Territory is the first in this region and
an important link between West Siberia and East
Siberia in the geographic distribution of S. kirchber-
gensis.
Irkutsk Province
A cranium and a mandible are known. The
locality for a mandible (IRM 2436) stored in the
Irkutsk local history museum (Table 1) is unknown.
It may belong to the same individual as a cranium
(ZIN 10718; Chersky, 1874), but the locality for the
cranium is also undefined.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
6
Institutional Abbreviations
IAE CHU: Institute of Archaeology and Ethnogra-
phy, Siberian Branch, Novosibirsk, collection from
Chumysh; IAE KY: Institute of Archaeology and
Ethnography, Siberian Branch, Novosibirsk, collec-
tion from Krasniy Yar (Novosibirsk Province); IAE
TRD: Institute of Archaeology and Ethnography,
Siberian Branch, Novosibirsk, collection from Tara-
danovo; IAE BB: Institute of Archaeology and Eth-
nography, Siberian Branch, Novosibirsk, collection
from Biysk; NSMLL: Novosibirsk State Museum of
Local Lore, Novosibirsk; IRM: Irkutsk Regional
Museum, Irkutsk; TRM: Tomsk Regional Museum,
Tomsk; KB MAN K: Kargasok branch of the
Museum of Art of the North, Kargasok, Tomsk
Province; PM TSU: Paleontological Museum of
Tomsk State University, Tomsk; Sk_oms1: Omsk
Province, the Bondarev private collection; Sk_ui1:
Omsk, the Bondarev private collection; GR PC:
The I. Grebnev private collection.
METHODS
Morphometrics
Descriptions of morphological and morpho-
metric features of teeth and measurements of
mandibles and postcranial bones were made fol-
lowing Gromova (1935), Guérin (1980), Lacombat
(2005, 2006), Kahlke (1977), Fortelius (1982),
Clauss et al. (2008), Dubrovo (1957), and Fortelius
et al. (1993).
The following dimensions were used for mor-
phometric analysis of rhinoceros mandibles:
1) overall length of mandible;
2) length from the mesial edge of the p2 alveoli
to the distal edge of the vertical ramus;
3) alveolar length of dentition p2-m3 (AL);
4) alveolar length of premolar row p2-p4;
5) alveolar length of molar row m1-m3
(ALm1_m3);
6) height of mandibular corpus between p2 and
p3 (HMB_p3);
7) height of mandibular corpus between p3 and
p4 (HMB_p4);
8) height of mandibular corpus between p4 and
m1 (HMB_m1);
9) height of mandibular corpus between m1 and
m2 (HMB_m2);
10) height of mandibular corpus between m2 and
m3 (HMB_m3);
11) transverse diameter of mandibular corpus
between p2 and p3 (TMB_p3);
12) transverse diameter of mandibular corpus
between p3 and p4 (TMB_p4);
13) transverse diameter of mandibular corpus
between p4 and m1 (TMB_m1);
14) transverse diameter of mandibular corpus
between m1 and m2 (TMB_m2);
15) transverse diameter of mandibular corpus
between m2 and m3 (TMB_m3);
16) length of mandibular symphysis;
17) length of diastema;
18) width of incisal area of the symphysis;
19) thickness of the symphysis distally;
20) transverse diameter of mandibular condyle;
21) height of mandibular condyle;
22) height from the level of the occlusal surface of
dentition to the mandibular condyle;
23) distance from the level of the mandibular
condyle to the distal edge of the m3 alveoli;
and
24) height of mandibular corpus before p2
(HMB_p2).
The following indices were used for compara-
tive analysis of Stephanorhinus kirchbergensis and
Coelodonta antiquitatis mandibles: index 1 =
ALm1_m3 / AL; index 2 = HMB_p2 / AL; index 3 =
HMB_p3 / AL; index 4 = HMB_p4 / AL; index 5 =
HMB_m1 / AL; index 6 = HMB_m2 / AL; index 7 =
HMB_m3 / AL; index 8 = TMB_p3 / AL; index 9 =
TMB_p4 / AL; index 10 = TMB_m1 / AL; index 11 =
TMB_m2 / AL; index 12 = TMB_m3 / AL.
The biological age of an individual animal was
determined based on the mandible by using the
method introduced by Hitchins (1978) and Hillman-
Smith et al. (1986). First, several stages reflecting
the degree of tooth development and wear were
identified.
The stages for upper teeth (STU) are:
1) presence of a dentine nucleus in the alveolus
(STU1);
2) dental crown erupted over the bone tissue but
unworn (STU2);
3) apices of most protruding cusps slightly worn
(STU3);
4) apices of most protruding cusps worn, but
wear does not affect steep ridges (STU4);
5) dental crown more worn than STU4, occlusal
surface has continuous sections (STU5);
6) dental crown worn by at least 50%, occlusal
part smooth, median valley still open (STU6);
7) dental crown worn more than 50%, median
valley closed (STU7);
PALAEO-ELECTRONICA.ORG
7
8) dental crown heavily worn, the median valley
closed by dentine on every side (STU8);
9) presence of few remains of enamel, fragmen-
tary mesial and distal valleys (median valley
disappeared) (STU9); and
10) absence of enamel, crown totally worn
(STU10).
The stages for lower teeth (STL) are:
1) erupting below the bone surface (STL1);
2) erupting above but unworn (STL2);
3) very slight wear, tips of cusps shiny (STL3);
4) slight wear exposing dentine on cusps, but
ridges steep with deep gaps (STL4);
5) more wear than STL4, but still with deep gaps
(STL5);
6) medium wear, tooth surface becoming fairly
flat, but infundibulum channel still open
(STL6);
7) medium/heavy wear, channel just closed
(STL7);
8) heavy wear, channel fully closed, through
dentine (STL8);
9) very heavy wear, small patches of enamel
remaining (STL9); and
10) no enamel remaining (STL10).
It is assumed that each tooth at each moment
of its biological age corresponded to one of the
above stages. Thus, the individual age of the ani-
mal could be defined by the cumulative information
on each tooth stage from the jaw tested, comparing
it to the stages given in the tables by Hillman-Smith
et al. (1986).
The following measurements were used to
analyze postcranial bones of rhinoceroses:
1) maximum length in the sagittal plane (ML);
2) antero-posterior diameter of the proximal
epiphysis (APD);
3) transverse diameter of the proximal epiphysis
(TD);
4) antero-posterior diameter of the distal epiphy-
sis (APDde);
5) transverse diameter of the distal epiphysis
(TDde);
6) transverse diameter of the distal joint (TDdj);
7) transverse diameter of the diaphysis in the
middle (mTDd);
8) antero-posterior diameter of the diaphysis in
the middle (mAPDd);
9) transverse diameter of the tuberositas tibia
(TDtt);
10) maximum transverse diameter measured per-
pendicularly to the vertical axis of the astraga-
lus (ATD);
11) maximum height, measured perpendicularly
to the first diameter of the astragalus (AH);
12) transverse diameter of distal joint of the
astragalus (ATD artic. dist.);
13) transverse diameter of distal part of the
astragalus below the collar (ATD max dist.);
14) transverse diameter of the olecranon (TD
olecr.);
15) antero-posterior diameter of the olecranon
(APD olecr.);
16) transverse diameter of the proximal joint (TD
artic. prox.); and
17) maximum height of the proximal joint (H artic.
prox.).
Mesowear Analysis
The mesowear method developed by Fortel-
ius and Solounias (2000) was applied to recon-
struct the dietary preferences of Stephanorhinus
kirchbergensis and Coelodonta antiquitatis in the
southeast area of West Siberia. The method is
based on the study of facet development on the
buccal side of occlusal surfaces of upper molar M2
of ungulates. The main factor in the formation of
facets in the area of the paracone and metacone is
the close interaction of the upper and lower teeth
during mastication (attrition). The other factor that
affects the formation of the occlusal surface is the
mechanichal properties of the food items that
crushed by the teeth. Consuming rough and abra-
sive food leads to smoothing of the wear surface of
the cusps by contact with each other. Mesowear
analysis deals with three variables: hypsodonty
index, wear surface relief (l, low; h, high), and the
shape of cusps (s, sharp; r, rounded; b, blunt). A
conservative approach was used to select the
paracone or metacone cusp (Fortelius and
Solounias, 2000), i.e., the sharpest one. The cusp
was considered high if the proportion of its height
to the crown width was larger than 0.03, which was
the empirical definition for rhinoceroses (Fortelius
and Solounias, 2000). Teeth in the first and last
dental wear stages were not considered in the
analysis. The method was applied for teeth in
stages STU5 to STU8. A data set of more than 20
specimens is considered statistically significant,
while a set of more than 10 individuals allows see-
ing the main vector of the adaptation process
toward one or another dietary preference.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
8
The mesowear method was extended by Kai-
ser and Solounias (2003) where not only M2 but all
possible combinations of upper teeth were used in
investigations of diet signals. In particular, the
research showed that statistical data obtained from
the set of four teeth Р4–М3 were sufficiently close
to those that were obtained while using only M2.
Thus, limited fossil material can be used by apply-
ing mesowear analysis to the set consisting of Р4–
М3.
Ten upper teeth of Stephanorhinus kirchber-
gensis presented in this article and two upper P4
and M1 from the Tomsk Province presented by
Shpansky and Billia (2012) were used as material
for mesowear analysis.
Statistical analysis was carried out using Sta-
tistical Package for the Social Sciences (SPSS)
version 21 and Sysstat version 12. Because the
independent samples of Stephanorhinus kirchber-
gensis and Coelodonta antiquitatis investigated
were not normally distributed, the nonparametric
Mann-Whitney U-test (SPSS 21) was used to
determine the degrees of difference of their
mesowear signals. Hierarchical cluster analysis
with complete linkage in a mode (the farthest
neighbor, with interval measure) of Euclidean dis-
tance was applied following the standard hierarchi-
cal amalgamation method. As an input for cluster
analysis, the percentages of independent
mesowear signals were used: the percentage of
high relief, the percentage of sharp peaks, and the
percentage of blunt peaks. Mesowear signals of C.
antiquitatis teeth from southern localities of West
Siberia from the collection of IAE were used as a
reference material. Data from subgroups of 27 typi-
cal recent animals presented in Fortelius and
Solounias (2000) were also used. Reference mate-
rial on Stephanorhinus was taken from Kahlke and
Kaiser (2011) and van Asperen and Kahlke (2014).
A conservative model of dietary classification was
used in classifying the animals by their dietary pref-
erences (Fortelius and Solounias, 2000).
Biogeochemistry
The study was conducted as part of an inter-
disciplinary research project on the study of bone
detritus of Cenozoic vertebrata in the Institute of
Geology, Komi Science Centre, Project Manager V.
I. Silaev. Analysis of the chemical and stable isoto-
pic composition was conducted using a broad set
of methods including several types of spectroscopy
and mass spectrometry. Methodological
approaches are described in more detail in a sepa-
rate paper (Silaev et al., 2015).
DESCRIPTION
Material of Stephanorhinus kirchbergensis
from 13 localities, consisting of 39 samples, is
described (Table 1). Published materials on the
European S. kirchbergensis and Coelodonta antiq-
uitatis were taken as the comparison.
Mandibles
Specimen NSMLL-12 (Figure 2, Table 2) is a
fragment of left mandibular corpus of a juvenile
individual.The rostral part is broken on the level of
symphysis, while the caudal part is absent behind
the alveolus for m1. The symphysis area shows
gnaw marks from small animals. The jaw is rela-
tively low; the width of the corpus gradually
increases from dp1 toward dp4. Widening of the
ventral part of the corpus is not visible. The mandi-
ble has an oval shape in the transverse plane,
while it is pear-shaped in Coelodonta antiquitatis.
Teeth dp2–4 are present. The metalophid of dp2 is
slightly worn, and the hypolophid is worn by 5–10%
(STL5); metalophid and hypolophid in dp3 are worn
for approximately 5–10% (STL5); dp4 is on the
fourth stage of wear (STL4). Alveolar length of the
tooth row dp1–dp4 is 138 mm. dp1 is absent. Its
alveolus is well developed and round. The alveolar
cavity for m1 is large. The edges of the alveolus
are directed toward each other, which shows the
initial stage of tooth eruption (formation of the
crown is at the first stage, STL1). The enamel is
smooth, without a clear pattern of enamel prisms.
The paraconid is absent in dp2. A fold-like undevel-
oped third valley near the parastylid is present on
dp3. The main valleys on dp2, dp3, and dp4 are
broad and U-shaped. Mesiodistal compression of
the crowns is not seen. Deciduous crowns have a
vertically convex shape on the buccal side. The
valley between lophids is deep. Lophids are con-
vex in occlusal view. In the first phase of the chew-
ing process, a browser forms the facets on the
buccal side of the lower teeth at an acute angle to
the occlusal surface. On the teeth of the given
mandible, these facets are in the initial stage of
forming. The underdeveloped folds of cingulid are
present on the lingual side of dp4 and in dp2–dp4
on the buccal side in the ventral part of the metalo-
phid. Cementum is absent. The estimated biologi-
cal age of the individual is less than two years.
All of the mandibles listed below for adult indi-
viduals show the following features. The mandibu-
lar corpus is relatively low. Its changes in height
and transversal diameter from p2 to m3 are linear;
the transverse section of the mandibular body is
oval, without ventral extension. Teeth are large with
PALAEO-ELECTRONICA.ORG
9
smooth enamel, without a clear pattern of enamel
prisms. All crowns are vertically convex and barrel-
shaped on the buccal side. The valley between the
lophids is distinct. The lophids of p3–m3 have a
convex surface in occlusal view. On the lower
teeth, there are facets on the buccal side at an
acute angle to the occlusal surface. These facets
were formed on the first phase of the browser’s
chewing process.
Specimen NSMLL-10 (Figure 3, Tables 3, 4) is
a fragment of mandible, the right mandibular body
of an adult individual. The symphysis is preserved
but is much damaged. The incisor edge of the sym-
physis is narrow, its dorsal surface is spoon-
shaped, and the ventral surface is convex. The
alveoli of the incisors are absent on the rostral
edge of the symphysis. The caudal edge of the
symphysis is on the level of p3, and the bone of the
mandible at the caudal side of the symphysis is
very thick. The left mandibular body is broken
between the alveoli of p3 and p4. The alveoli of p2
and p3 contain roots, but the crowns are
destroyed. The right mandibular corpus is pre-
served together with the angular area, while the
vertical ramus is broken off. The right dentary row
p2–m3 is completely preserved. The diastema is
relatively short. Dentition is highly worn: p2 (STL9),
p3 (STL9), p4 (STL8), m1 (STL8), m2 (STL7), and
m3 (STL6). The valleys are V-, or U-shaped and
preserved only on m2 and m3. In p2–m2, there is a
poorly developed cingulid on the buccal side of the
ventral part of the metalophid and on the lingual
side of the paraconid. Remains of a very thin layer
of cementum are preserved on the back wall of m2
on the root area of the crown. A considerable level
of wear suggests a biological age of the animal
close to 30 years.
Specimen NSMLL 22090 (Figure 4, Tables 3,
4) is fragment of mandible, the left mandibular cor-
pus of an adult individual. The rostral part is broken
FIGURE 2. Stephanorhinus kirchbergensis; Chumysh
River at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia); the horizontal left body of a
mandible of a juvenile rhino NSMLL-12, occlusal and
buccal views. Scale bar ruled in centimeters.
TABLE 2. metric comparison of lower deciduous teeth of stephanorhinus kirchbergensis and coelodonta antiquitatis
of the middle and upper pleistocene from the altay territory and novosibirsk and tomsk provinces in southeastern
western siberia versus european and chinese regions. all measurements are in mm. sample sizes are given in
parentheses. Available in zipped download at https://palaeo-electronica.org/content/2021/3322-stephanorhinus-
kirchbergensis-in-siberia.
TABLE 3. Metric comparison of lower permanent teeth of Stephanorhinus kirchbergensis and Coelodonta antiquita-
tis of the Middle and Upper Pleistocene from the Altay Territory and Novosibirsk and Tomsk Provinces in southeast-
ern Western Siberia versus European and Chinese areas. All measurements are in mm. Sample sizes are given in
parentheses. Available in zipped download at https://palaeo-electronica.org/content/2021/3322-stephanorhinus-
kirchbergensis-in-siberia.
TABLE 4. Metric comparison of the mandibles of Stephanorhinus kirchbergensis and Coelodonta antiquitatis of the
Middle and Upper Pleistocene from the Altay Territory and Novosibirsk, Tomsk, Omsk, and Irkutsk Provinces of
southeastern Western Siberia versus European and Chinese areas. All measurements are in mm. Sample sizes are
given in parentheses. Available in zipped download at https://palaeo-electronica.org/content/2021/3322-stepha-
norhinus-kirchbergensis-in-siberia.
FIGURE 3. Stephanorhinus kirchbergensis; Chumysh
River at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia); the horizontal right body of
a mandible NSMLL-10, occlusal and buccal views.
Scale bar ruled in centimeters.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
10
near the p2, while the caudal end is lacking the
portion behind the alveolus of m3. All teeth are
present in the jaw from p2 to m3. They have an
average stage of wear: p2 (STL7), p3 (STL7), p4
(STL6), m1 (STL7) m2 (STL6), and m3 (STL4).
The trignoid basin of p4, m1, and m2 is wide and V-
shaped. The mesial and distal valleys of m3 are
wide and U-shaped. A poorly developed cingulid is
present on the buccal side of the ventral part
metalophid in p4, m2, and m3, and on the lingual
side from the paraconid to the metaconid in p2–
m2. Cementum is absent. The individual’s esti-
mated biological age is 16–20 years.
Specimen IAE KY-4323 (Figure 5, Tables 3, 4)
is a fragment of mandible, the left mandibular cor-
pus of an adult individual. The rostral part is broken
near p2. The caudal part is broken in the angular
area. All teeth p2–m3 are present in the jaw but
heavily worn: p2 (STL9), p3 (STL9), p4 (STL9), m1
(STL9), m2 (STL8), and m3 (STL7). The trigonid
basin of m3 is U-shaped. The cingulid is present on
p2–m2 on the buccal side of the ventral part of the
metalophid and on the lingual side of the paraconid
towards the metaconid. The cementum is not pre-
served. This is the smallest of all specimens pre-
sented in this overview, which may be correlated
with sexual dimorphism and suggest an identifica-
tion as a female individual. The estimated biologi-
cal age of the individual is more than 30 years.
Specimen Sk_ui1 (Figure 6, Tables 3, 4), is a
fragment of a mandible, the right mandibular cor-
pus of an adult individual. The specimen is broken
off near the front edge of the p2 alveolus and the
back edge of the m3 alveolus. The symphysis is
partially preserved and is high and long, with an
abrupt caudal edge at the middle part of p3. The
tooth row from p3 to m3 is preserved in the jaw and
shows average stages of wear: p3 (STL7), p4
(STL6), m1 (STL7), m2 (STL6), and m3 (STL4).
The trigonid basin of m2 is V-shaped. Both valleys
of m3 are wide and U-shaped. Folds of the cingu-
lum are observed in the basal part of m2 and m3
crowns on the buccal side of the hypolophid and on
the metalophids of p3 and p4. Signs of a thin layer
of cementum are preserved on m1–m3. The esti-
mated age of the individual is 20–25 years.
Specimen GR PC 1164 (Figure 7, Tables 3, 4)
is a fragment of mandible, right horizontal mandib-
ular corpus of adult individual. The symphysis is
broken off near the alveolus of p2. The coronoid
FIGURE 4. Stephanorhinus kirchbergensis; Ob River at
Krasniy Yar (Novosibirsk Province, southeast Western
Siberia); the horizontal left body of a mandible NSMLL
22090, occlusal and buccal views. Scale bar ruled in
centimeters.
FIGURE 5. Stephanorhinus kirchbergensis; Ob River at
Krasniy Yar (Novosibirsk Province, southeast Western
Siberia); the horizontal left body of a mandible IAE KY-
4323, occlusal and buccal views. Scale bar ruled in cen-
timeters.
FIGURE 6. Stephanorhinus kirchbergensis; Omsk
(Omsk Province, southeast Western Siberia); the hori-
zontal right body of a mandible Sk_ui1, occlusal and
buccal views. Scale bar ruled in centimeters.
PALAEO-ELECTRONICA.ORG
11
process is broken off from the vertical ramus. The
mandibular condyle is broken from the lateral side.
The mandibular corpus is very low compared to the
other specimens. The notch of the symphysis is
placed on the level of p3. The vertical ramus is rel-
atively broad with a strongly developed masseteric
fossa. The angle between the vertical ramus and
the body is close to 90º. The mandibular body pre-
serves p3–m1 but crowns are damaged. The
crowns of p3 and p4 are worn to the level of the
bottom of the mesial valley (STL7). Distal valleys
are V-shaped. The m1 is worn to the level of the
bottom of the distal valley (STL8). The signs of cin-
gulid are observed on p3 on the lophid buccal side,
and on the lingual side on p4 and m1. The distal
edge of the m3 alveolus is close to the base of the
vertical ramus. The area behind the m3 alveolus is
wide and flat, with rounded edges. The estimated
biological age of the individual is 20–25 years.
Specimen GR PC 1165 (Figure 8, Tables 3, 4)
is the well-preserved mandible of adult individual.
There is some damage to the right p2 alveolus and
right coronoid process. The incisors’ alveoli are
absent. The left m1 (STL9) and m2 (STL8) are pre-
served but significantly damaged; the rest of the
dentition is broken or missing. The right dentition is
preserved, showing the high degree of wear: p4
(STL8), m1 (STL9), and m2 (STL8). Well-devel-
oped р2 alveoli show that there was no natural loss
and overgrowth of alveoli in this specimen. The
diastema is short. The notch of symphysis is
placed on the level of p3, the bone of mandible at
the caudal extremity of symphysis is very thick.
The symphysis is long and narrow, while the dias-
tema is relatively short. Its dorsal surface is spoon-
shaped. Its ventral surface is convex. The dorsal
area behind the m3 is wide and flat with rounded
edges. The vertical ramus of the mandible is rela-
tively wide with strongly developed masseteric
fossa. The angle between the vertical ramus and
corpus is close to 90°. The left coronoid process is
directed upwards; its tip does not bend in the cau-
dal direction, as in Coelodonta antiquitatis. The
condyles are very wide with a visible asymmetry in
lateral direction. The medial edge of the condyle
has a clear rectangular shape in caudal view. The
lateral edge of the caudal part of the area posterior
to the condyle process rises in a smooth, concave
curve towards the condyle. In C. antiquitatis, this
edge approaching the condyle becomes rounded
towards the vertical axis of the vertical ramus of the
mandible and becomes convex. The estimated bio-
logical age of the individual is 25–30 years.
Specimen IRM 2436 (Figure 9, Table 4) is a
mandible of an adult individual with broken sym-
physis and coronoid processes. The notch of sym-
physis is placed at the level of p3, the bone at the
FIGURE 7. Stephanorhinus kirchbergensis; Biysk (Altay
Territory, southeast Western Siberia); the horizontal right
body of a mandible GR PC-1164, occlusal and buccal
views. Scale bar ruled in centimeters.
FIGURE 8. Stephanorhinus kirchbergensis; Biysk
(Altay Territory, southeast Western Siberia); the mandi-
ble GR PC-1165, 1, occlusal view. 2, posterior view. 3,
buccal view. Scale bar ruled in centimeters.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
12
caudal side of symphysis is very thick. The dorsal
area behind the m3 is wide and flat with rounded
edges. The mandibular body is relatively low. Its
changes in height and transversal diameter from
p2 to m3 are linear. The transverse section of the
corpus is oval, without ventral extension. The verti-
cal ramus is relatively broad with a strongly devel-
oped masseteric fossa. The angle between the
vertical ramus and horizontal body is close to 90°.
The condyles are very wide with a visible lateral
asymmetry. The dentition is partly absent; the rest
of it is broken. The paraconid, parastylid, and
metaconid are broken on the left p4. The enamel is
mostly absent; therefore the presence of cingulid is
unclear. Mesial and distal valleys are shallow
(depth of the distal valley is 7.0 mm). The thickness
of the enamel on the hypoconid is 2.0 mm on the
buccal side, 1.4 mm on the medial side. The
metaconid is slightly worn. The estimated age of
the individual is 20–25 years.
Specimen KB MAN K-397 (Figure 10.3-10.5,
Tables 3, 4) is a fragment of mandible, left mandib-
ular corpus of an adult individual, whose biological
age is estimated to 15–17 years. The p3-m3 teeth
are preserved in the jaw. The caudal edge of the
symphysis is on the same level as the middle part
of p3, similarly to the condition seen in the speci-
men from Cherniy Yar (Gromova, 1935). The ven-
tral edge of the mandibular body is almost smooth.
The rise of the ventral edge is gradual from m1 to
the symphysis, with increasing roundness in the
symphysis area. Ventral margin is not concave in
Stephanorhinus kirchbergensis. The thickness of
the mandibular body is nearly the same along the
whole tooth row, slightly decreasing under p3-p4.
In transverse section, the mandibular corpus is
high and oval-shaped, while it is pear-shaped with
a ventral bulge in adult Сoelodonta antiquitatis.
Mental foramens (one large and several small) are
placed under the p2 alveolus. The vertical ramus is
relatively wide, with well-developed rugosity in the
FIGURE 9. Stephanorhinus kirchbergensis; Irkutsk
Province (southwest Eastern Siberia); a mandible IRM
2436, occlusal and buccal views. Scale bar ruled in
centimeters.
FIGURE 10. Stephanorhinus kirchbergensis; Tobol Hori-
zon level (Middle Pleistocene); at Asino (Asino District,
Tomsk Province, southeast Western Siberia); first left
upper molar M1 PM TSU 1/396, 1. buccal view. 2, occlu-
sal view. KB MAN K-397 the horizontal left body of a
mandible of the Tobol Horizon level (Middle Pleisto-
cene); Ob River at Kindal village (Kargasok District,
Tomsk Province, southeast Western Siberia). 3, buccal
view. 4, occlusal view. 5, occlusal view р3 – m3.
PALAEO-ELECTRONICA.ORG
13
angular area. The dorsal surface of the mandibular
body behind the m3 is wide (62 mm), flattened,
with a small longitudinal groove; the edges of the
area are not sharp, as mentioned by Gromova
(1935), and gently rounded, and the ridges are not
developed. Its back surface forms an area that is
broad and slightly concave in lateral view, perpen-
dicular to the longitudinal axis of the jaw. In woolly
rhinoceros, this area is considerably smaller, trian-
gular, and oriented by an significant angle to the
longitudinal axis. The articular head is inclined rela-
tive to the horizontal plane. Its medial edge is
absent, and the lateral one is raised. Teeth crowns
are high. The m3 is in the first stage of wear. Pre-
molars are placed vertically to the alveolar edge of
the mandibular body. Molars are significantly
inclined forwards (Figure 10.3). Teeth are large,
especially m1. Metrically, they are close to the
samples from Moldavia, Povolzhy, and Taubach,
but smaller than teeth from Krasniy Yar (Tomsk
region). The length of the molar row m1–3 is 171
mm, which is larger than that in European samples.
The closest length is in a large sample from Tau-
bach (157.8–169.9 mm; Kahlke, 1977). Jaws from
Chumysh River (NSMLL-10) and Kraskiy Yar
(Novosibirsk Province; NSMLL 22090) (Table 4)
show similar dental size. The buccal cingulid is well
defined on all metalophids and hypolophids of m1
and m2. The lingual cingulid is well developed on
the metalophids of m1 and m2, and on the base of
m3. The crown width near the roots is larger than
on the apex, because of the slight inclination of the
buccal surface. It is especially well observed on the
slightly worn m2 and m3. In C. antiquitatis, the
crown width is constant along the height. In this
specimen of S. kirchbergensis, the metalophid on
the molars is shorter than the hypolophid. The
walls of lophids expand greatly to the base, forming
valleys highly narrowed sharply downward.
Metaconids are tapered roundly (without bulbosi-
ties) and are slightly abducted backwards on the
protoconids of m2 and m3. The cementum frag-
ments are preserved in the basal part. The enamel
is smooth. There is a fusion of metalophid and
hypolophid in p3–m1, the valleys are considerably
deep. The fusion of metalophid and hypolophid on
m2 and m3 has not occurred (Figure 10.5). The
mesial surface of metaconid is unworn on m3.
Specimen NSMLL-26 (Figure 11.3-4, Table 3)
is a fragment of mandible, right mandibular body of
an adult individual. The rostral part is broken off at
the level of the symphysis notch, between the p2
and p3 alveoli. The caudal side is broken off
between the p4 and m1 alveoli. The alveolus of p4
bears the remains of roots. The p3 (STL6) is pre-
served in the jaw. Valleys are V-shaped. A weakly
developed cingulid is present on the buccal and lin-
gual sides of hypolophid. The estimated biological
age of the individual is 6–9 years.
Specimen NSMLL-27 (Figure 11.1-2, Table 3)
is a fragment of mandible, right vertical ramus of an
adult individual. The vertical ramus is relatively
broad with a strongly developed area of attach-
ment of the masseteric muscles to the masseteric
fossa. The coronoid process is directed upwards.
Its tip bends in the aboral direction, as it is fre-
quently observed in Coelodonta antiquitatis. The
condyle is very wide (transverse diameter is 112
mm) with a visible lateral asymmetry. The medial
edge of the condyle is sharp, with a rectangular
outline. In C. antiquitatis, this edge becomes
rounded towards the vertical axis of the vertical
ramus when approaching the condyle and it takes
a convex shape.
Isolated Teeth
Upper teeth are large, brachyodont, with
smooth enamel, and without a clear pattern of
FIGURE 11. Stephanorhinus kirchbergensis; Chumysh
River at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia); the vertical right ramus of
a mandible NSMLL-26. 1, posterior view. 2, lingual
view. The horizontal right body of a mandible NSMLL-
27. 3, buccal viw. 4, occlusal view. Scale bar ruled in
centimeters.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
14
enamel prisms. Medial valleys are open, except in
two M3 (NSMLL-4 and IAE BB-1), for which they
are closed. The occlusal surface of the teeth is
concave, except for the specimen PM TSU 1/396
that displays a flat occlusal surface.
The upper left М1 PM TSU 1/396 (Figure
10.1-2, Table 5) is well preserved, with some
enamel breakages on the parastyle and metastyle.
The roots are preserved. The tooth is similar to PM
TSU 5/3495 from Krasniy Yar (Shpansky and Billia,
2012) in the degree of wear and morphology. The
protoloph and hypocone are bulbous. The front
wall of the protoloph bears part of a cingulum. The
enamel is smooth, with a porcelain gloss. The
antecrochet is absent, the crista is poorly devel-
oped, and the crochet is well developed and quad-
rate-shaped.
Specimen NSMLL-5 (Figure 12.1, Table 5) is
an upper left M2 that has an average stage of wear
(STU6). Crochet is well developed, without virga-
tions. Crista and antecrochet are absent. The cin-
gulum is clear, present on the mesial part of the
tooth, and only slightly developed on the lingual
side. The protocone is clearly separated. The style
of the paracone is well developed from the side of
ectoloph. The buccal side of the tooth is signifi-
cantly higher than the lingual side. The ectoloph is
inclined in the lingual direction and serrated. The
relief of the ectoloph is high. Apices of the
paracone and metacone are rounded. Fragments
of a thin layer of cementum are present.
Specimen NSMLL-1 (Figure 12.2, Table 5) is
an upper right M2 that is moderately worn (STU5).
The crochet is well developed, without virgations.
The crista is well developed, with double virgation.
Antecrochet is absent. The cingulum is present on
the mesial part of the tooth, while it is slightly visi-
ble on the lingual part of the tooth. The protocone
is not isolated. The style of the paracone is well
developed from the side of the ectoloph. The buc-
cal side of the tooth is significantly higher than the
lingual side. The ectoloph is inclined in the lingual
direction and serrated. The relief of the ectoloph is
high. The apices of the paracone and metacone
are rounded.
Specimen NSMLL-6 (Figure 13.1, Table 5) is
an upper left P4 that is more than 50% worn
(STU8). The crochet is well developed, without vir-
gations. The crista is absent. Antecrochet is
absent. A poorly-defined cingulum is observed on
the lingual part of the tooth. The protocone is not
isolated. The occlusal surface is concave. The
relief of the ectoloph is high. Apices of the
paracone and metacone are rounded.
TABLE 5. Comparison of the upper permanent teeth of Stephanorhinus kirchbergensis and Coelodonta antiquitatis of
the Middle and Upper Pleistocene from the Altay Territory and Novosibirsk, and Tomsk Provinces of southeastern
Western Siberia versus European and Chinese areas. All measurements are in mm. Sample sizes are given in paren-
theses. Available in zipped download at https://palaeo-electronica.org/content/2021/3322-stephanorhinus-kirchbergen-
sis-in-siberia.
FIGURE 12. Stephanorhinus kirchbergensis; Chumysh
River, at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia). 1, second left upper molar
M2, NSMLL-5. 2, second right upper molar M2, NSMLL-
1; occlusal and anterior views. Scale bar ruled in centi-
meters.
FIGURE 13. Stephanorhinus kirchbergensis; Chumysh
River, at Kytmanovo (Kytmanovo District, Altay Terri-
tory, southeast Western Siberia). 1, fourth left upper
permanent premolar P4, NSMLL-6. 2, fourth right upper
permanent premolar P4, NSMLL-2; occlusal and ante-
rior views. Scale bar ruled in centimeters.
PALAEO-ELECTRONICA.ORG
15
Specimen NSMLL-2 (Figure 13.2, Table 5) is
an upper right P4 that is more than 50% worn
(STU8), with partially destroyed protocone and
parastyle. Crochet is well developed, without virga-
tions. Crista and antecrochet are absent. A poorly-
defined cingulum is observed on the lingual part of
the tooth. The protocone is not isolated. The occlu-
sal surface is concave. The relief of the ectoloph is
high. The apex of the metacone is sharp. Frag-
ments of a thin layer of cementum are present.
Specimen NSMLL-7 (Figure 14.1, Table 5) is
an upper left P4 that is 50% worn (STU7). Crochet
is well developed, with double virgation. The crista
is well developed, without virgations. Antecrochet
is absent. Well-developed cingula are present on
the mesial and lingual sides of the tooth. The proto-
cone is not isolated. The relief of the ectoloph is
high. The apices of the paracone and metacone
are sharp.
Specimen NSMLL-8 (Figure 14.2, Table 5) is
an upper left P4 worn more than 50% (STU8). The
crochet is well developed, and unlike the crista,
without virgations. Antecrochet is absent. Well-
developed cingula are observed on the mesial and
lingual sides of the tooth. The protocone is not iso-
lated. The relief of the ectoloph is high. The apices
of the paracone and metacone are rounded.
Specimen NSMLL-3 (Figure 15.2, Table 5) is
an upper left M3 that is moderately worn (STU6),
with destroyed protocone and parastyle. The cro-
chet is well developed, without virgations. The sin-
gle crista is poorly developed, without virgations.
Antecrochet is absent. The tooth is triangular with
merged ecto- and metaloph in the dorsal section.
The relief of the occlusal surface is high. The api-
ces of the paracone and metacone are destroyed.
Specimen NSMLL 21052 (Figure 15.1, Table
5) is an upper right M2 that is more than 50% worn
(STU8). The ectoloph is preserved as a narrow
stripe because of the high degree of wear but it is
FIGURE 14. Stephanorhinus kirchbergensis; Chumysh
River, at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia). 1, fourth left upper perma-
nent premolar P4, NSMLL-7. 2, fourth left upper perma-
nent premolar P4, NSMLL-8; occlusal and anterior
views. Scale bar ruled in centimeters.
FIGURE 15. Stephanorhinus kirchbergensis; 1, Ob
River (Novosibirsk Province, southeast Western Sibe-
ria); second right upper molar M2, NSMLL 21052,
occlusal and anterior views. 2, Chumysh River, at Kyt-
manovo (Kytmanovo District, Altay Territory, southeast
Western Siberia); third right upper molar M3, NSMLL-3,
occlusal and buccal views. 3, third left upper molar M3,
NSMLL-4, occlusal view. 4, third left upper molar M3,
IAE BB-1, Berdsk (Novosibirsk Province, southeast
Western Siberia), occlusal and buccal views. 5 and 6.
Omsk (Omsk Province, southeast Western Siberia),
third left lower premolar p4, Sk_oms1. 5, buccal view. 6,
occlusal view. Scale bar ruled in centimeters.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
16
serrated. The relief of the ectoloph is high. The api-
ces of the paracone and metacone are sharp. The
cingulum is poorly developed on the lingual side.
The protocone is not isolated.
Specimen NSMLL-4 (Figure 15.4, Table 5) is
an upper left M3 that is moderately worn (STU5).
The roots of the tooth are broken. The crochet and
crista are well developed, without virgations. Ante-
crochet is absent. The median valley is closed. The
cingulum is present only on the mesial side. The
protocone is not isolated. The style of the paracone
is well developed from the side of the ectoloph.
The relief of the ectoloph is high. The apices of the
paracone and metacone are rounded. Fragments
of a thin layer of cementum are present. The tooth
is triangular with merged ecto- and metaloph in the
dorsal section.
Specimen IAE BB-1 (Figure 15.3, Table 5) is
an upper left molar M3 that is moderately worn
(STU6). The crochet and crista are well developed,
without virgations. Antecrochet is absent. The
median valley is closed. A well-defined cingulum is
present on the mesial side of the tooth. The proto-
cone is not isolated. The style of the paracone is
well developed from the side of ectoloph. The relief
of the ectoloph is high. The apices of the paracone
and metacone are rounded. Fragments of a thin
layer of cementum are present. The tooth is trian-
gular with merged ecto- and metaloph in the dorsal
section.
Specimen Sk_oms1. (Figure 15.5-6, Table 3)
is a lower left p4. The tooth is large and heavily
worn to the level of the distal valley. The crown is
vertically convex and barrel-shaped. The valley
between lophids is well defined. The lophids are
convex on the occlusal surface. There is a facet on
the buccal side, which forms an acute angle to the
occlusal surface. This facet appears in rhinocer-
oses that are characterized by a brachyodont den-
tition.
The lower left р3 PM TSU 1/395 (Table 3) is a
tooth that is heavily worn below the level of valleys.
The enamel is preserved only on the buccal wall of
the hypolophid. The enamel is light gray, smooth,
and porcelaneous, without wrinkles or cementum.
The front edge of the metalophid has a horizontal
cylinder-shaped bulbosity at the base. The metalo-
phid is narrower than the hypolophid.
Postcranial material
Specimen NSMLL-109 (Figure 16.1-3, Table
6) is a well-preserved left ulna that lacks the distal
epiphysis. The bone is long and gracile. Its abso-
lute size is smaller than the equivalent bone from
the Taubach (Kahlke, 1977). In Coelodonta antiqui-
tatis, the mediolateral diameter of some individuals
is considerably larger than that. In the given speci-
men, the proc. coracoideus slightly rises upwards
(Figure 16.1), which is a characteristic of S. kirch-
bergensis. In C. antiquitatis the proc. coracoideus
points downwards and the proximal part of the
olecranon has a wide flat area, which abruptly
expands towards the olecranon tubercle on the
medial side. This expansion forms a sharp edge,
pointed medially. In NSMLL-109, the proximal part
of the olecranon is relatively narrow with a sharp-
edged ridge along the dorsal surface, and a
smooth extension on the medial side towards the
olecranon tubercle not forming the medial ridge
(Figure 16.2). In C. antiquitatis, a notch extends
from outside of the ventro-medial projection of inci-
sura semilunaris, and around its distal portion,
which sharply separates it from the rest of the joint
surface of the diaphysis. In NSMLL-109, this notch
is not observed. The transition between the joint
and the diaphysis is smooth (Figure 16.3). There is
a ridge along the lateral side of the diaphysis from
the incisura semilunaris to the distal epiphysis with
a gap in the middle. In the area of the gap, the
diaphysis is relatively sharp and forms a dihedral
angle with a rounded tip (Figure 16.2). In C. antiq-
uitatis, that area of the diaphysis is flat. In NSMLL-
109, the palmar wall of the diaphysis is straight
and, unlike C. antiquitatis, without flexure.
Specimen IAE CHU-108 (Figure 17.1-4, Table
7) is a well-preserved right radius of Stephanorhi-
nus kirchbergensis. It has a large absolute length
of 436 mm; it is larger than similar bones of Coe-
lodonta antiquitatis (maximum length for the speci-
men from Krasniy Yar, Tomsk Province, is 424 mm)
(Shpansky, 2014). Specimen IAE CHU-108
appears more slender, the transverse diameter of
the proximal epiphysis index being 25.2% of the
total length of the bone, while the average values
are 26.2% and 29.7% for S. kirchbergensis and C.
antiquitatis, respectively (Guérin, 1980). Absolute
values for the measurements and indices of the rel-
ative proportions correspond to the values typical
for S. kirchbergensis. In C. antiquitatis, the transi-
tion from the ventral edge of the lateral part of the
proximal epiphysis facet to the lateral edge is
smooth, rounded, and convex and the intersection
of the lateral and dorsal edges forms an obtuse
angle of around 120°. In IAE CHU-108, the ventral
edge of the proximal facet of the lateral part is
straight and considerably shifted towards the
medial facet, which makes the ventral edge appear
shorter. The transition in the lateral edge occurs by
PALAEO-ELECTRONICA.ORG
17
FIGURE 16. Stephanorhinus kirchbergensis; Chumysh River at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia). The left ulna, NSMLL-109. 1, medial view. 2, anterior view. 3, lateral view. The right
astragalus, NSMLL-107. 4, dorsal view. 5, plantar view. Scale bar ruled in centimeters.
TABLE 6. Measurements of the ulna of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeastern
Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following mea-
surements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiphysis
(APD); antero-posterior diameter of the distal epiphysis (APDde); transverse diameter of the distal epiphysis (TDde);
transverse diameter of the diaphysis in the middle (mTDd); antero-posterior diameter of the diaphysis in the middle
(mAPDd); transverse diameter of the olecranon (TD olecr.); antero-posterior diameter of the olecranon (APD olecr.);
transverse diameter of the proximal joint (TD artic. prox.); maximum height of the proximal joint (H artic. prox.).
Dimensions
S. kirchbergensis C. antiquitatis
Altay Territory,
NSMLL-109
Europe, Taubach
(Kahlke, 1977)
Europe (Guérin, 1980 tab.
131) Europe (Guérin, 1980 tab. 131)
Range Mean Range Mean
ML 456 - 543 494.5(21)
TD olecr. 72 67 – 68 67.5(2) 57 - 102 84.2(13)
APD olecr. 90 103 - 107 104.5(3) 90 - 120 106.4(23)
TD artic.prox. 98 101.8 91 – 94 92.5(2) 75 - 109 92.5(30)
APD 175 140 - 157 148.5(2) 121 - 195 159.1(26)
mTDd 43 49.9 51 – 52 51.5(2) 44 – 68 55.4(18)
mAPDd 41 46.3 52 52(2) 41.5 - 60 51.8(18)
TDde 40 41 – 67 53.8(16)
APDde 77 60 – 92 76.2(15)
H artic. prox. 113 106.5
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
18
a rounded obtuse angle, around 130°. The lateral
edge is straight and intersects a dorsal edge at an
acute rounded angle, less than 90° (Figure 17.3).
This morphology of the radius proximal joint is typi-
cal of S. kirchbergensis (Guérin, 1980). In C. antiq-
uitatis, the tuberosity on the dorsal surface of the
distal epiphysis is divided by a broad, shallow sag-
ittal groove. In IAE CHU-108, the tuberosity on the
dorsal surface of the distal epiphysis is flat, without
a visible separation (Figure 17.1), which is charac-
teristic of S. kirchbergensis (Guérin, 1980). In C.
antiquitatis, the dorsal ridge separating the facets
for lunate and scaphoid bones on the surface of
the distal joint is well defined, especially on the
anterior part of the joint surface (Gromova, 1950).
In IAE CHU-108, the dorsal ridge is almost invisible
(Figure 17.4).
Specimens NSMLL-102 and GR PC 203 are
well-preserved left Mc II and GR PC 214 is a right
Mc II that is damaged in the area of the distal joint
and near the facet for Mc III (Figures 18.1-4, 19.1-
8, Table 8). All of the bones listed above are large
(Table 8) and considerably exceed the dimensions
of the Coelodonta antiquitatis bones. Moreover, the
bones of Stephanorhinus kirchbergensis are more
slender. Their three main transverse diameter indi-
ces are considerably smaller than those in C. antiq-
uitatis. The surface of the Мс II proximal joint from
the lateral side has a sharp angle between trape-
zoid and magnum facets. In C. antiquitatis, it forms
an angle of around 100°, while in NSMLL-102 and
GR PC 203, it is less than 90° (Figures 18.3, 19.3).
The lateral facet for Mc III in C. antiquitatis has a
more abrupt expansion than in S. kirchbergensis
(Guérin, 1980) (Figures 18.2, 19.2). The cross-sec-
tion of the diaphysis is more elliptic with a small
keel along the volar surface, especially in GR PC
214 (Figure 19.7). The diaphysis is thicker from the
medial side, which characterizes S. kirchbergensis
(Guérin, 1980). In C. antiquitatis, the cross-section
of the diaphysis has a broadly elliptic shape with a
relatively sharp edge on the medial side and the
surface of the diaphysis on the volar side is either
flat or tuberous (Guérin, 1980).
Specimens IAE TRD-17 and IAE TRD-18 (Fig-
ures 18.5-8, 20.1-4, Table 9) are left Mc IV. Speci-
men IAE TRD-18 has lengthwise damage in the
area of the distal epiphysis, while specimen IAE
TRD-17 is well preserved. The bones have a large
relative length and are considerably larger than
those in Coelodonta antiquitatis. At the same time,
the bones are more slender; their epiphysis and
diaphysis indices are much smaller than those in
C. antiquitatis (Table 9). In C. antiquitatis, the volar
side of the proximal joint (the closest distance
between the facet for Mc V and volar facet for Mc
III) is always smaller than the dorsal side of the
joint (the closest distance between the facet for Mc
V and dorsal facet for Mc III). Both dorsal and volar
sides of the joint in these specimens have approxi-
mately the same length, just as in Stephanorhinus
kirchbergensis (Guérin, 1980). In C. antiquitatis,
the sides of the two facets for articulation with Mc
III are usually fused in the medial part of the proxi-
mal epiphysis, forming a dihedral angle, but may
be separated by a small groove. In S. kirchbergen-
sis, the groove is present between the facets. In C.
antiquitatis, the palmar facet for Mc III is usually
round while the dorsal palmar facet is relatively
wider (the height is more than half the width). In
IAE TRD-17 and IAE TRD-18, the palmar facet is
FIGURE 17. Stephanorhinus kirchbergensis; Chumysh
River at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia). The right radius, IAE CHU-
108. 1, anterior view. 2, lateral view. 3, proximal view. 4,
distal view. The left tibia, NSMLL-110. 5, distal view. 6,
proximal view. 7, anterior view. 8, posterior view. Scale
bar ruled in centimeters.
PALAEO-ELECTRONICA.ORG
19
high and relatively narrow, while the dorsal one is
low and long (the width is about a third of the
height), with a small inclination towards the proxi-
mal edge (Figures 18.7, 20.3), as mentioned for S.
kirchbergensis (Guérin, 1980). In IAE TRD-17 and
IAE TRD-18, the tuberosity for attachment of the
interosseous muscle on the medial surface is rela-
tively weak (Figures 18.5, 20.1), which is not typi-
cal of C. antiquitatis (Belyaeva, 1966). In C.
antiquitatis, the cross-section of the diaphysis is
relatively thick and triangular, with rounded angles.
In S. kirchbergensis, just as in IAE TRD-17 and
IAE TRD-18, the cross-section of the Mc IV diaphy-
sis is trapezoidal in shape with a large base on the
dorsal side and a very acute angle at the base of
the lateral side (Guérin, 1980).
Specimen PM TSU 5/5197 is a Mc III with a
strongly flattened diaphysis in the dorsoventral
direction. The distal end of the bone is absent (Fig-
ure 21.3-4). The bone is large (Table 10), signifi-
cantly exceeding the previously described
specimen PM TSU 5/2723 (Figure 21.1-2) (Shpan-
sky and Billia, 2012), but smaller than a massive
bone from Rybinsk (Belyaeva, 1940).
Specimen NSMLL-110 (Figure 17.5-8, Table
11) consists of a left tibia and a left fibula. The
bones are fused and well preserved. The fibula has
the following sizes: absolute length is 351 mm (in
Coelodonta tologoiensis, 340 mm [Belyaeva,
1966]); its mediolateral width of the proximal epiph-
ysis is 28 mm; its transverse width of the proximal
epiphysis is 44 mm; its mediolateral width of the
diaphysis is 32 mm; its transverse diameter of the
distal end is 54 mm; and its transverse width index
is 15.4% (in Coelodonta antiquitatis, it is less than
16% [Gromova, 1950]). Tibia length of NSMLL-110
is 444 mm, which exceeds the size of C. antiquita-
tis (maximal length for the Krasniy Yar locality of
Tomsk Province, West Siberia, is 438 mm) (Shpan-
sky, 2014). The tibia of S. kirchbergensis appears
more slender than that of C. antiquitatis (Table 11;
indices of proximal and distal epiphyses). Absolute
values for measurements of tibia NSMLL-110 and
indices of relative proportions correspond to the
values typical of S. kirchbergensis. Tuberositas on
the proximal epiphysis of tibia is relatively short
and less massive (maximum mediolateral width of
tuberositas to width of the proximal epiphysis is
36%). Therefore, the width of the proximal epiphy-
sis is significantly larger than the transverse diame-
ter (Figure 17.6), which is a characteristic of
Stephanorhinus kirchbergensis. In C. antiquitatis,
maximum width of tuberositas is about 50% of the
width of the proximal epiphysis, which makes the
width of the proximal epiphysis less than its trans-
verse diameter. In S. kirchbergensis, and in
NSMLL-110, the medial edge of the tibia distal
TABLE 7. Measurements of the radius of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeast-
ern Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following
measurements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiph-
ysis (APD); transverse diameter of the proximal epiphysis (TD); antero-posterior diameter of the distal epiphysis
(APDde); transverse diameter of the distal epiphysis (TDde); transverse diameter of the diaphysis in the middle
(mTDd).
Dimensions
S. kirchbergensis C. antiquitatis
Altay Territory,
IAE CHU-108
Europe
(Guérin, 1980 tab. 130)
Europe
(Guérin, 1980 tab. 130)
Altay Territory and
Novosibirsk Province, IAE:
Krasniy Yar, Taradanovo,
Chumysh
Range Mean Range Mean Range Mean
ML 436 408 - 445 421.8(5) 334 - 413 380.4(81) 335 – 405 379.1(13)
APD 88 68 - 87 74.9(18) 55 - 93 77.5(106) 68.5 – 91.5 81.6(13)
TD 110 102 - 119 110.4(18) 97 - 126 112.8(109) 95 - 123.5 114.7(13)
APDde 66 61 - 82 67.9(10) 62 - 92 76.6(80) 61 - 84.5 76.1(13)
TDde 109 90.5 - 113.5 105(10) 95 - 142 117.7(84) 94 - 130.5 118.6(13)
mTDd 57 53 - 65 58.7(9) 54 - 75.5 63.4(103) 52.7 – 73 63.7(13)
Indexes, %
TD/ML 25.2 26.2(5) 29.7(81) 28.4 - 32.3 30.3(13)
mTDd/ML 13.1 13.9(5) 16.7(81) 15.4 – 18 16.8(13)
TDde/ML 25 24.9(5) 30.9(81) 28.1 - 32.5 31.3(13)
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
20
epiphysis is convex (Figure 17.5), while in C. antiq-
uitatis, it is concave (Gromova, 1950).
Specimen NSMLL-107 (Figure 16.4-5, Table
12) is a right astragalus. The bone is large, close to
the size of the specimen from Krasniy Yar (Tomsk
Province) and Western Europe (Table 11). The
ratio of the height of the bone to its width is 93.3%;
the transverse diameter compared to the width is
58.5%. The average values for Stephanorhinus
kirchbergensis are 94.2% and 65.9%, respectively;
for Coelodonta antiquitatis, they are 90.8% and
65.2%, respectively (Guérin, 1980). The absolute
values for the measurements and proportional indi-
ces for S. kirchbergensis usually exceed the values
for C. antiquitatis, but some large individuals of
woolly rhinoceros may display similar sizes. In S.
kirchbergensis, just as in NSMLL-110, the dorsal
part of the lateral calcaneal facet has a sharp pos-
terior edge without transition into the ventral part
(Figure 16.5), as in C. antiquitatis (Shpansky and
Billia, 2012).
Specimens PM TSU 5/2538 and PM TSU 5/
3063 are right naviculars of adult individuals (Fig-
ure 22). Because the bones of the postcranial skel-
eton of Stephanorhinus kirchbergensis can often
be found together with the remains of a woolly rhi-
noceros in alluvial localities during redeposition, we
compared the characteristics and features of the
navicular bone of these species (Table 13).
Specimen NSMLL-101 (Figure 23.1-4, Table
14) is a left Mt II. The specimen is slightly worn in
the area of the proximal epiphysis and has a large
absolute length. The bone appears slender; its pro-
portions are close to the average of Stephanorhi-
nus kirchbergensis from Europe (Guérin, 1980).
Two facets conjoined with facets of Mt III are pres-
ent on the lateral side of the proximal epiphysis.
The dorsal facet is smaller than the plantar one, its
contour is near trapezoid in shape, and the proxi-
mal edge does not exceed the proximal edge of the
plantar facet. The plantar facet is trapezoid-
shaped; its height is larger than its width. Both fac-
ets have an unclear outline, which is typical for S.
kirchbergensis (Guérin, 1980). In Coelodonta
antiquitatis, the two facets for Mt III are usually
placed on the lateral side of the proximal epiphysis,
but often they may be merged. If the facets are
separate, the dorsal one may be either round or
elliptical and proximo-distally extended. The plan-
tar facet is usually round. The dorsal facet is
placed higher than the plantar one. There is a facet
for the first cuneiform on the plantar side of the
proximal epiphysis. In S. kirchbergensis, the facet
is relatively large, usually rectangular or the shape
of a flipped L (Guérin, 1980). In C. antiquitatis, this
facet is placed high, elliptical or trapezoid, and
often limited by the deep vertical groove. In
NSMLL-101, the plantar facet is unclear because
of the damage on this spot; the vertical groove is
absent. The diaphysis of S. kirchbergensis is pen-
tagonal in the dorsal plane, while four front angles
may have an unclear outline, but on the plantar
side there is always a clear V-shaped angle, due to
the high keel along the middle part of plantar wall
surface. This keel is clearly observed on NSMLL-
110 (Figure 23.3), but the distal expansion of the
diaphysis is relatively weaker than in other species
of rhinoceroses (Guérin, 1980). In C. antiquitatis,
the diaphysis is elliptical in the dorsal plane and
more circular and its distal expansion is larger,
compared to the other species of rhinoceroses
(Guérin, 1980).
Specimens NSMLL-105 and IAE TRD-2 (Fig-
ures 20.5-8, 23.5-8, Table 15) are well-preserved
left Mt IV. The length of those specimens is higher
than in Coelodonta antiquitatis and Mt IV of the
European samples of Stephanorhinus kirchbergen-
sis (Guérin, 1980). The relative medio-lateral width
of the S. kirchbergensis diaphysis and the epiphy-
ses of Mt IV is less than those of C. antiquitatis
(Table 15). In S. kirchbergensis, part of the dorsal
edge of proximal is concave from the dorsal facet
for Mt III in the lateral direction as the dorsal facet
for Mt III stretches dramatically in the medio-plan-
tar direction; we get a distinct change in the outline
of the dorsal edge of the proximal facet, which has
an angle of about 100°. This characteristic angle is
well defined on the proximal facets of NSMLL-105
and IAE TRD-2 (Figures 20.8, 23.8). In C. antiqui-
tatis, this area of the proximal facet dorsal edge is
convex without sharp bends. At the same time, the
outline of the proximal facet plantar edge is highly
concave in NSMLL-105 and IAE TRD-2 (Figures
20.8, 23.8), which is typical of S. kirchbergensis
and not typical for C. antiquitatis (Belyaeva, 1966).
A calloused protuberance is well developed on the
plantar side of the proximal epiphysis in NSMLL-
105 and IAE TRD-2. In NSMLL-105, it rises in the
proximal direction to the level of the joint surface
(Figure 23.8). This is not observed on the proximal
epiphysis of C. antiquitatis. In NSMLL-105 and IAE
TRD-2, the dorsal facet is semi-elliptical and
touches the edge of the proximal facet. Its width is
less than half of the length. The plantar facet has
the shape of an ellipse that is slightly elongated
vertically; it is placed below the dorsal facet and
separated from the proximal facet by a notch (Fig-
ures 20.6, 23.6), similar to S. kirchbergensis
PALAEO-ELECTRONICA.ORG
21
(Guérin, 1980). In C. antiquitatis, if both facets are
present, the dorsal one is semicircular, with its
width little larger than its height, and the plantar
facet is circular. When the facets are united, the
common facet is L-shaped (Guérin, 1980). In S.
kirchbergensis, just as in NSMLL-105 and IAE
TRD-2, the cross-section of the diaphysis is oval
and stretched in the dorso-plantar direction without
sharp inflections in the contour. In C. antiquitatis,
the diaphysis has a round cross-section with sharp
bends due to the frequent presence of sharp longi-
tudinal ridges on the surface of the diaphysis
(Guérin, 1980).
ODONTOLOGICAL ANALYSIS
Adaptation of some animal groups to a food
base that included a high proportion of abrasive
food (grass) led to significant physiological and
morphological differences from those who pre-
ferred a diet with a small percentage of rough food
(Clauss et al., 2008). Recent and Pleistocene rhi-
noceroses are one of the most prominent repre-
sentatives that demonstrate a high degree of
specialization of adaptation to different ecological
environments. Therefore, considering the morpho-
logical features of the fossil material represented
by Stephanorhinus kirchbergensis, we focused on
the morphological characteristics that reflected
food preferences. For greater clarity, the study was
carried out by comparing S. kirchbergensis with the
morphology of Coelodonta antiquitatis that lived in
West Siberia. We used the methods presented in
Fortelius (1982) and Clauss et al. (2008) to study
the morphological differences of typical browsers
and grazers based on the recent African rhinocer-
oses Diceros bicornis and Ceratotherium simum.
First of all, let us consider the adaptations
associated with increased resistance of the teeth to
abrasion caused by different dietary requirements.
The considerable amount of silica in the grass and
its cover of dust in the Pleistocene landscapes
under arid climate significantly affected the rate of
dental wear. Therefore, the appearance of hypsod-
FIGURE 18. Stephanorhinus kirchbergensis; Chumysh River at Kytmanovo (Kytmanovo District, Altay Territory, south-
east Western Siberia). The left Mc II, NSMLL-102. 1, dorsal view. 2, lateral view. 3, volar view. 4, proximal view. Ob
River at Taradanovo Village (Suzun District, Novosibirsk Province, southeast Western Siberia). The left Mc IV, IAE
TRD-17. 5, dorsal view. 6, medial view. 7, volar view. 8, proximal view. Scale bar ruled in centimeters.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
22
FIGURE 19. Stephanorhinus kirchbergensis; Bijsk (Altay Territori, southeast Western Siberia). The left Mc II, GR PC-
203. 1, dorsal view. 2, lateral view. 3, volar view. 4, proximal view. The right Mc II, GR PC-214, 5. dorsal view. 6, lateral
view. 7, volar view. 8, proximal view. Scale bar ruled in centimeters.
TABLE 8. Measurements of McII of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeastern
Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following mea-
surements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiphysis
(APD); transverse diameter of the proximal epiphysis (TD); antero-posterior diameter of the distal epiphysis (APDde);
transverse diameter of the distal epiphysis (TDde); transverse diameter of the diaphysis in the middle (mTDd).
Dimensions
S. kirchbergensis C. antiquitatis
Altay
Territory
NSMLL-
102
Altay
Territory
GR PC-
203
Altay
Territory
GR PC-
214
Europe
(Guérin, 1980 tab. 139)
Europe
(Guérin, 1980 tab.
139)
Altay Territory and
Novosibirsk Province,
IAE: Krasniy Yar,
Taradanovo, Chumysh
Range Mean Range Mean Range Mean
ML 187.5 186 c194 179 – 212 195.4(13) 148 - 180 164.2(60) 144.7 - 180.4 164.3(26)
APD 46 62.5 59 42.5 – 60 48.2(12) 40 - 58 47(57) 35 – 52 44.7(26)
TD 48 58.5 61 41 – 57 48.0(12) 41 - 66.5 52.9(61) 47 – 63 54.8(26)
APDde 44 51.5 37.5 - 53.7 45.0(12) 35 - 52.5 43.1(58) 35 – 49 42.8(26)
TDde 52 52 57 45 – 56 48.9(12) 37.5 - 57 48.8(55) 41.4 - 56.5 47.7(26)
mTDd 40 40 44 33.5 - 41.5 38.6(13) 31.5 - 50 42.4(60) 37 - 48.7 42(26)
Indexes, %
TD/ML 25.6 31.5 31.4 24.6(12) 32.2(60) 29.6 - 36.2 33.4(26)
mTDd/ML 21.3 21.5 22.7 19.7(13) 25.8(60) 23.8 - 28.7 25.5(26)
TDde/ML 27.7 28 29.4 25(12) 29.7(55) 26.2 – 32 29(26)
PALAEO-ELECTRONICA.ORG
23
FIGURE 20. Stephanorhinus kirchbergensis; Ob River at Taradanovo village (Suzun District, Novosibirsk Province,
southeast Western Siberia). The left Mc IV, IAE TRD-18. 1, dorsal view. 2, medial view. 3, volar view. 4, proximal view.
The left Mt IV, IAE TRD-2. 5, dorsal view. 6, medial view. 7, plantar view. 8, proximal view. Scale bar ruled in centime-
ters.
TABLE 9. Measurements of McIV of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeastern
Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following mea-
surements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiphysis
(APD); transverse diameter of the proximal epiphysis (TD); antero-posterior diameter of the distal epiphysis (APDde);
transverse diameter of the distal epiphysis (TDde); transverse diameter of the diaphysis in the middle (mTDd).
Dimensions
S. kirchbergensis C. antiquitatis
Novosibirsk
Province,
IAE TRD-17
Novosibirsk
Province,
IAE TRD-18
Europe
(Guérin, 1980 tab. 141)
Europe
(Guérin, 1980 tab. 141)
Altay Territory and
Novosibirsk Province,
IAE: Krasniy Yar,
Taradanovo, Chumysh
Range Mean Range Mean Range Mean
ML 199 188.5 172.5 - 193 182.1(9) 126.5 - 176.5 151.1(59) 136 - 163.5 151.3(25)
APD 52 51.5 39 - 51 43.3(12) 39 – 52 45(52) 35 – 49 44.9(25)
TD 55.5 51 48 - 62 51.6(12) 41 - 62.5 53.3(57) 47 – 61 55.4(25)
APDde 51 0 41 - 50 44.9(9) 34 – 48 42.3(50) 37 – 45 41.5(25)
TDde 56 52.5 43 - 51 47.3(8) 42 - 62.5 47.9(58) 43 – 55 49.6(25)
mTDd 44.5 40 34 - 42 38.1(10) 32 – 46 37.6(59) 30.5 – 48 38.6(25)
Indexes, %
TD/ML 27.9 27 28.3(9) 35.3(57) 34.6 – 38.4 36.6(25)
mTDd/ML 22.4 21.2 20.9(9) 24.9(59) 22.4 - 29.4 25.5(25)
TDde/ML 28.1 27.9 26(8) 31.7(58) 31.2 - 35.6 32.8(25)
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
24
onty, as a result of adaptation to increased wear of
the teeth, is observed in many groups of animals.
We use the description of the morphological differ-
ences between hypsodont and brachyodont types
of teeth in rhinos (Fortelius, 1982). The upper teeth
described here show the main features of the
brachyodont type, different from the characteristics
of the hypsodont type observed in Coelodonta
antiquitatis.
The occlusal surface of the upper teeth of S.
kirchbergensis is concave (Figure 24.3); in Coe-
lodonta antiquitatis, the occlusal surface is flat (Fig-
ure 24.4). Up to STU7, the buccal side of the upper
teeth of S. kirchbergensis is higher than the lingual
and highly inclined to the lingual side, i.e., there is
a bucco-lingual constriction in the occlusal area of
the crown in slightly worn teeth (Figure 12.1-2;
mesial view). Teeth of this type are characterized
by uneven wear of the occlusal surface. After
STU7, the crown height on the buccal and lingual
sides becomes equal (Figure 14.1), and the buccal
side becomes shorter than the lingual in the follow-
ing stages (Figures 13.1-2, 14.2, 15.1). In C. antiq-
uitatis, almost vertical buccal and lingual walls
have nearly equal height, and wear is equal. In S.
kirchbergensis, the ectoloph of all upper teeth is
serrated. In C. antiquitatis, the ectoloph is straight.
In S. kirchbergensis, secondary folds of all upper
premolars and molars do not merge with each
other until the very last stages of wear and do not
form closed inner valleys. In C. antiquitatis, the
secondary folds usually merge in the very first
stages of wear, forming the closed inner valleys. In
some specimens of S. kirchbergensis, styles are
FIGURE 21. Stephanorhinus kirchbergensis; Tobol Hori-
zon level (Middle Pleistocene), Ob River at Krasniy Yar
(Tomsk Province, southeast Western Siberia). The right
Mc III, PM TSU 5/5197. 1, lateral view. 2, dorsal view.
The left Mc III, PM TSU 5/2723. 3, lateral view. 4, dorsal
view. Scale bar ruled in centimeters.
TAB LE 10 . Measurements of McIII of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeastern
Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following mea-
surements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiphysis
(APD); transverse diameter of the proximal epiphysis (TD); antero-posterior diameter of the distal epiphysis (APDde);
transverse diameter of the distal epiphysis (TDde); transverse diameter of the diaphysis in the middle (mTDd); trans-
verse diameter of the distal joint (TDdj).
Dimensions
S. kirchbergensis C .antiquitatis
Tom sk
Province,
Krasniy
Yar,
PM TSU 5/
5197
Krasniy Yar
(Shpansky
and Billia,
2012)
PM TSU 5/
2723
Rybinsk
(Beljaev
a, 1939)
Taubach
(Kalke,
1977)
Western Europe:
(Guérin, 1980, tab. 140)
Tom sk
Province,
Krasniy
Yar (QIII)
PM TSU
Western Europe:
(Guérin, 1980, tab.
140)
Range Mean n=7 Range Mean
ML С186 229 225 204.2 206-250.5 225.2(13) 164-198 162-213 189(79)
APD С50 56.6 64 63.7-67.8 50-59 54.2(13) 42-58 42.5-61.5 52.2(80)
TD 70 63 80 63.7-67.8 58-71 63.9(17) 58.2-79 59.5-79 68.2(90)
APDde – 56 64 48-58.5 52.8(14) 41.8-55 44-57.5 50.8(68)
TDde – 80.4 90 73.9 64.5-83 73.8(14) 55-71 57.5-74 65.8(77)
TDdj – 64 70 59.1 52-64.5 59.6(13) – 49-65 56.1(77)
mTDd 67.5 60.5 72 53.5-55.7 54-70.5 61.4(16) 48-58 46-66 56.4(86)
Indexes, %
TD/ ML – 27.5 35.6 31.2 28.4 33-39.9 36.1(79)
TDde/ ML – 35.1 40 36.2 32.8 31.9-36.2 34.8(77)
mTDd/ ML – 26.4 31.1 26.2 27.3 28.2-30.1 29.9(79)
PALAEO-ELECTRONICA.ORG
25
TAB LE 11. Measurements of tibia of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeastern
Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following mea-
surements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiphysis
(APD); transverse diameter of the proximal epiphysis (TD); antero-posterior diameter of the distal epiphysis (APDde);
transverse diameter of the distal epiphysis (TDde); transverse diameter of the diaphysis in the middle (mTDd); trans-
verse diameter of the diaphysis in the middle (mTDd); antero-posterior diameter of the diaphysis in the middle
(mAPDd); transverse diameter of the tuberositas tibia (TDtt).
Dimensions
S. kirchbergensis C. antiquitatis
Altay
Ter rit ory,
NSMLL-110
Europe
(Guérin, 1980 tab. 144)
Europe
(Guérin, 1980 tab. 144)
Altay Territory and Novosibirsk
Province IAE: Krasniy Yar,
Taradanovo, Chumysh
Range Mean Range Mean Range Mean
ML 444 404 - 457 429(3) 323.5 - 433 381.1(67) 364 - 424 394.5(4)
APD 133 137.5(1) 87 - 157.5 136.8(42) 125 - 156 141.3(4)
TD 149 136(1) 111 - 163 133.4(50) 119 - 147 134.4(4)
APDde 90 75.5 - 92 85.6(12) 70 - 98 82.3(88) 77 - 91.5 85.3(4)
TDde 122.5 105 - 128 111.8(12) 92 - 127 106.8(88) 101 - 120 109.4(4)
mAPDd 62 60.5 - 69.5 64.4(5) 51 - 77 63.2(82) 57 - 67 64(4)
mTDd 69 63.5 - 80 70.5(5) 59 - 82.5 70.1(85) 65 - 73 69.3(4)
TDtt 54 60 - 74 66.8(4)
Indexes, %
TD/ML 33.6 31.7(1) 35(50) 32.3 - 36.5 34(4)
mTDd/ML 14 16.4(3) 18.4(67) 15.3 - 17.3 16.2(4)
TDde/ML 27.6 26(3) 28(67) 26.2 - 28.6 27.7(4)
APD/ML 30 32.1(1) 35.9(42) 34.1 - 37.9 35.8(4)
mAPDd/ML 15.5 15(3) 16.6(67) 17.2 - 17.9 17.6(4)
APDde/ML 20.3 20(3) 21.6(67) 20.2 - 23.6 21.6(4)
TDtt/TD 36.2 43.8 - 54.6 49.8(4)
TABLE 12. Measurements of the astragalus of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from south-
eastern Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The follow-
ing measurements were used: maximum transverse diameter measured perpendicularly to the vertical axis of the
astragalus (ATD); maximum height, measured perpendicularly to the first diameter of the astragalus (AH); transverse
diameter of distal joint of the astragalus (ATD artic. dist.); transverse diameter of distal part of the astragalus below the
collar (ATD max dist.).
Dimensions
S. kirchbergensis C. antiquitatis
AltayTerritory,
Tomsk
Province,
Krasniy Yar
(Shpansky and
Billia, 2012)
Europe (Guérin, 1980
tab. 145)
Tomsk
Province,
Krasniy Yar
(Shpansky and
Billia, 2012)
Europe (Guérin, 1980
tab. 145)
NSMLL-107 PM TSU 5/740 Range Mean n=39 Range Mean
ATD 112 113 93 – 113 101.7(31) 82-111 84 - 112 95.7(112)
AH 104.5 104 85 – 105 95.8(29) 79-98 77 - 102 87(112)
ATD artic. dist. 93 93 74 – 93 84.7(29) 68 – 91 80.9(107)
ATD maxi dist. 97.5 96 79 – 99 89(30) 85-98 75 – 97 85.1(108)
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
26
isolated. For example, isolation of the protocone is
observed (Figure 12.1; occlusal view). In C. antiq-
uitatis, styles are not isolated. In S. kirchbergensis,
the thickness of enamel is uneven along the perim-
eter of the tooth. It is thicker on the buccal and lin-
gual sides. In C. antiquitatis, the thickness of the
enamel is even along the tooth perimeter. In S.
kirchbergensis, the shape of M3 in the dorsal plane
is close to triangular, with merged ecto- and
metalophs (Figure 15.2-4). In C. antiquitatis, the
M3 cross-section is quadrate in shape, and the
metaloph is often separated from the ectoloph. In
all lower teeth of S. kirchbergensis, the lophids are
convex on the occlusal side. In C. antiquitatis,
lophids are flattened on the occlusal side. All lower
teeth of S. kirchbergensis are convex on the buccal
side, i.e., there is a smooth expansion from the
occlusal part of the crown and a smooth constric-
tion to the root. In C. antiquitatis, the walls on the
buccal and lingual sides are almost vertical and
flattened. All of the upper and lower teeth of S.
kirchbergensis are characterized by relatively
smooth enamel without a clear pattern of enamel
prisms, by the absence of rugosity, and by the rare
presence of cementum. In C. antiquitatis, the
enamel surface is rough, generally with a distinct
pattern of enamel prisms or wrinkling. Cementum
is retained and often covers a large portion of the
tooth crown in C. antiquitatis.
Further, it is necessary to review the results of
adaptations associated with different ways of grind-
ing the food for brachyodont and hypsodont types
of teeth. The chewing process of brachyodont
teeth is in two phases (Fortelius, 1982; Popowics
and Fortelius, 1997; Steuer et al., 2010). In the first
phase (the cutting phase), the sharp edge of the
ectoloph interacts only with the dorsal buccal edge
of the lower teeth, producing the primary tearing of
FIGURE 22. The navicular: Stephanorhinus kirchber-
gensis, Tobol Horizon level (Middle Pleistocene), Ob
River at Krasniy Yar (Tomsk Province, southeast West-
ern Siberia). 1, PM TSU 5/2538. 2. PM TSU 5/3063.
Coelodonta antiquitatis, Karginian Horizon, Ob River at
Krasniy Yar (Tomsk Province, southeast Western Sibe-
ria), 3. PM TSU 5/3815. а, proximal view. b, distal view.
c, dorsal-medial view.
TAB LE 1 3. Morphological comparison of the navicular bone of Stephanorhinus kirchbergensis versus Coelodonta
antiquitatis.
Stephanorhinus kirchbergensis Coelodonta antiquitatis
1. The proximal articular surface has a wide notch in the
latero-plantar side.
The notch is either very weak or absent.
2. The dorso-lateral angle is not very acute and only slightly
elongated. Correlatively with this, the width of the
proximal articular surface is approximately the same as
the length.
The dorso-lateral angle always well pronounced and
markedly elongated in the lateral direction. The proximal
articular surface is more extended latero-medially.
3. The process talocaudalis (which is used for connection
with a cubic bone) is displaced to the plantar edge of the
lateral side.
The process talocaudalis is placed at the middle of the
lateral side.
4. The dorso-medial angle has a convex shape. The dorso-medial angle is oblique.
5. The joint, on the distal side, has a well pronounced
indentation in the proximal direction between facets for
connection with III cuneiform and I cuneiform.
This indentation is no such pronounced.
6. The III cuneiform facet has a well pronounced gap from
the dorso-lateral edge of the bone.
This gap is no such pronounced.
PALAEO-ELECTRONICA.ORG
27
the plant tissues. In this stage, a brief but intense
stress concentrated on a small area of surface is
produced. Only the serrated edge of the ectoloph
participates in this phase. The shape of the ectol-
oph curves follows the shape of curves in lophids
of the lower teeth and the dorsal buccal edge of
lophids in the lower teeth. This leads to the occur-
rence of polished facets on the buccal edge of
lophids, which are at an acute angle to the occlusal
surface (Figure 24.1). In the second phase (the
crushing phase), there is an interaction of the main
part of the occlusal surface for both upper and
lower teeth, with a translocation in the oral-aboral
direction (Figure 24.5). The features of this masti-
catory apparatus in Stephanorhinus kirchbergensis
were developed from a juvenile age. A characteris-
tic facet is observed on the milk teeth of the juve-
nile mandible NSMLL-12 (Figure 2). In
brachyodont teeth, both phases are well devel-
oped, while in hypsodont teeth, the first phase is
rudimentary and the second phase proves to be
the main one, representing the crushing under high
pressure and large side amplitude. As a result, the
occlusal surface of the upper and lower teeth of
Coelodonta antiquitatis is flat (Figure 24.2, 24.4,
24.6).
The next adaptations are caused by different
chewing forces for brachyodont and hypsodont
types of teeth. A large proportion of herbaceous
plants in the diet of Coelodonta antiquitatis
increases the force required for its milling. Mandi-
bles of large herbivores are forced to move laterally
with a large amplitude, when the animal is required
to create a maximum force to break the plant tis-
sue, which has in its structure long, longitudinal,
durable, and elastic fibers (as in grass). If the feed-
ing objects have a mosaic structure, as does the
leaf of a shrub or young tree, movements back and
forth are enough to break it (Sanson, 2006). The
increase in forces in herbivores is reached by rele-
vant morphological adaptations of the mandible. In
this case, the relative height and thickness of the
horizontal corpus and the relative height of the
mandibular condyle above the level of the tooth
row are increased. All of the mandibles studied
appear more slender than C. antiquitatis according
FIGURE 23. Stephanorhinus kirchbergensis; Chumysh River at Kytmanovo (Kytmanovo District, Altay Territory,
southeast Western Siberia). The left Mt II, NSMLL-101. 1, dorsal view. 2, lateral view. 3, plantar view. 4, proximal view.
The left Mt IV, NSMLL-105. 5, dorsal view. 6, medial view. 7, plantar view. 8, proximal view. Scale bar ruled in centime-
ters.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
28
to characteristics listed above. It can be seen on
Figure 25, where indices of molars relative length
(Figure 25; Ind. 1), horizontal corpus relative height
(Figure 25; Ind. 2-7) and thickness (Figure 25; Ind.
8-12) for S. kirchbergensis are lower than those for
C. antiquitatis.
The adaptation to a more selective manner of
feeding is presented by a very narrow, spoon-like
mandibular symphysis area and shortened viscero-
cranium, because of the reduced diastema (Table
4, Figure 3, Figure 8.1, occlusal view). In Coe-
lodonta antiquitatis, the area of mandibular sym-
TABLE 14. Measurements of MtII of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeastern
Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following mea-
surements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiphysis
(APD); transverse diameter of the proximal epiphysis (TD); antero-posterior diameter of the distal epiphysis (APDde);
transverse diameter of the distal epiphysis (TDde); transverse diameter of the diaphysis in the middle (mTDd).
TABLE 15. Measurements of MtIV of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from southeastern
Western Siberia and Europe. All measurements are in mm. Sample sizes are given in parentheses. The following mea-
surements were used: maximum length in the sagittal plane (ML); antero-posterior diameter of the proximal epiphysis
(APD); transverse diameter of the proximal epiphysis (TD); antero-posterior diameter of the distal epiphysis (APDde);
transverse diameter of the distal epiphysis (TDde); transverse diameter of the diaphysis in the middle (mTDd).
Dimensions
S. kirchbergensis C. antiquitatis
Altay Territory,
NSMLL-101
Europe (Guérin, 1980 tab.
152)
Europe (Guérin, 1980
tab. 152)
Altay Territory and
Novosibirsk Province, IAE:
Krasniy Yar, Taradanovo,
Chumysh
Range Mean Range Mean Range Mean
ML 188 173.5 - 195 180.7(7) 140 - 157.5 148.5(37) 130.5 - 176.8 146.9(16)
APD 43.5 44 - 51 47.1(9) 36.5 - 51 41.8(34) 37 - 51.6 43(16)
TD 34 31 - 39 34.8(9) 27.5 - 38 32.6(36) 25.6 – 39.3 32.6(16)
APDde 46 41 - 48.5 43.2(7) 33.5 - 43 38.1(33) 34.7 – 46.7 37.7(16)
TDde 41 38 - 44 41(6) 31.5 - 44.5 37.6(36) 30 - 47.5 36.4(16)
mTDd 31 26.5 - 33.5 29.1(7) 23.5 - 37 31.1(37) 25.5 – 33.8 29.3(16)
Indexes, %
TD/ML 18.1 19.2(7) 21.8(36) 17.7 – 27.1 22.3(16)
mTDd/ML 16.5 16.1(7) 20.9(37) 18.6 – 22.5 20(16)
TDde/ML 21.8 22.7(6) 25.3(36) 22.4 – 27.7 24.8(16)
Dimensions
S. kirchbergensis C. antiquitatis
Altay Territory,
NSMLL-105
Novosibirsk
Province,
IAE TRD-2
Europe (Guérin, 1980
tab. 154)
Europe (Guérin, 1980
tab. 154)
Altay Territory and
Novosibirsk Province,
IAE: Krasniy Yar,
Taradanovo, Chumysh
Range Mean Range Mean Range Mean
ML 188.5 176 170 - 182.5 178.2(3) 127 - 155 144.9(40) 122.2 – 163 146.8(15)
APD 54.5 51.5 44 - 53 47.2(6) 37 - 51.5 44.3(39) 37.1 - 51.5 46.3(15)
TD 54 53 47 - 53.5 50.1(6) 41 - 57 46.5(37) 36.7 - 52.5 46.4(15)
APDde 52 46.5 44.5 - 51.5 48.8(3) 36 - 46 40.8(35) 36.3 – 47 42.5(15)
TDde 43 40 37 - 43 40.3(4) 31 - 41 36.1(36) 29.1 – 45 36.1(15)
mTDd 35 28.5 33.5 - 36.5 34.8(4) 24 - 40 30.5(40) 22.2 – 41 30.6(15)
Indexes, %
TD/ML 28.6 30.1 28.1(3) 32.1(37) 27.1 - 37.5 31.7(15)
mTDd/ML 18.6 16.2 19.5(3) 21.1(40) 16.1 - 26.4 20.8(15)
TDde/ML 22.8 22.7 22.6(3) 24.9(36) 22.2 - 27.6 24.6(15)
PALAEO-ELECTRONICA.ORG
29
physis is flat, wide, and relatively long (Table 4).
Another result of this adaptation may be a less
wide opening of the mouth. In Stephanorhinus
kirchbergensis, this is caused by the more acute
angle of rise in the vertical ramus of the mandible
(Figures 7, 10). In C. antiquitatis, the vertical ramus
rises slightly, which allows the mouth to open
wider. Another adaptation to a more selective man-
ner of feeding could be the high body position and
slender structure of the S. kirchbergensis postcra-
niun. This is well illustrated by the morphometric
data of the material presented (Tables 6, 7, 8, 9,
10, 11, 12, 13, 14, 15, Figures 16, 17, 18, 19, 20,
21, 22, 23). The structure of the occipital region of
the cranium allowed feeding on the vegetation at a
higher level. Distinctive features of C. antiquitatis
are the adaptations for gathering food from the
ground: shortened limbs, a strongly overhanging
occipital crest that prevents lifting the head high,
and an elongated viscerocranium caused by the
shifting of orbits in the caudal direction.
MESOWEAR ANALYSIS
Characteristics of 12 teeth of Stephanorhinus
kirchbergensis were used as an input data for
mesowear analysis. Ten of them were character-
ized in the present work, and data about two teeth
(P4 and M1) were taken from the article of Shpan-
sky and Billia (2012) (Table 16).
All of the studied upper molars of Stepha-
norhinus kirchbergensis from West Siberia had a
high occlusal relief. A high percentage of teeth
FIGURE 24. The specifics of the wear of the occlusal
surfaces of the lower and upper teeth of Stephanorhinus
kirchbergensis (1,3,5) and Coelodonta antiquitatis
(2,4,6). Arrows indicate the morphological differences
associated with a different biomechanical masticatory
system for the browser and the grazer. Scale bar ruled in
centimeters.
FIGURE 25. Indexes of the mandible of Stephanorhinus kirchbergensis and Coelodonta antiquitatis.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
30
have sharp apices (41.6%). The rest of the teeth
have rounded apices, none of them have blunt api-
ces (Table 17). In C. antiquitatis from the same
region, only 28% had a high occlusal relief, 64% of
specimens had rounded cusps, and 36% had blunt
cusps; no specimen displayed sharp cusps (Table
17).
Even the conservative assessment of hypsod-
ont index (on lower third molars from the material
presented) gives values close to 1.7 for Stepha-
TABLE 16. Mesowear scores for occlusal relief and cusp shape for maxillary teeth of fossils Stephanorhinus kirchber-
gensis and Coelodonta antiquitatis.
Species locality
Specimen identification
number Upper tooth
occlusal relief
(high=h, low=l)
cusp shape
(sharp=s,
round=r, blunt=b)
Stephanorhinus
kirchbergensis
fossils from the
southeast of
Western Siberia
IAE CHU-1 M2 h R
IAE CHU-5 M2 h S
IAE CHU-2 P4 h S
IAE CHU-6 P4 h S
IAE CHU-7 P4 h S
IAE CHU-8 P4 h R
IAE BB-1 M3 h R
IAE CHU-4 M3 h R
PM TSU 5/3495 M1 h R
PM TSU 5/2878 P4 h R
PM TSU 5/396 M1 h R
NSMLL 21052 M2 h S
Coelodonta
antiquitatis
IAE CHU 51 P4 l B
IAE CHU 52 P4 l R
IAE CHU 53 P4 l R
IAE CHU 54 P4 l B
IAE CHU 55 P4 l R
IAE CHU 56 M1 l B
IAE CHU 57 M1 l R
IAE CHU 58 M1 l B
IAE CHU 59 M1 l B
IAE CHU 60 M1 l B
IAE CHU 61 M1 l R
IAE CHU 62 M2 l R
IAE CHU 63 M2 l R
IAE CHU 64 M2 h R
IAE CHU 65 M2 h R
IAE CHU 66 M2 l R
IAE CHU 67 M2 h R
IAE CHU 68 M2 l B
IAE CHU 69 M2 l R
IAE CHU 70 M2 l R
IAE CHU 71 M3 l B
IAE CHU 72 M3 h R
IAE CHU 73 M3 h R
IAE CHU 74 M3 h R
IAE CHU 75 M3 h R
PALAEO-ELECTRONICA.ORG
31
norhinus kirchbergensis from West Siberia and 2
or a little larger for Coelodonta antiquitatis. Thus,
S. kirchbergensis can be classified as brachyodont
type, while C. antiquitatis belongs to the mesodont
type. But, as shown in by Fortelius and Solounias
(2000), using only the hypsodont index does not
allow classifying definitely the individuals. In addi-
tion, this characteristic affects slightly the results of
mesowear analysis.
Two groups of attributes of dental specimens
were used as an input for a Mann-Whitney U-test:
one for Stephanorhinus kirchbergensis and the
other for Coelodonta antiquitatis. The values of
attributes were formed based on two mesowear
variables (relief of the occlusal surface and the
shape of cusps) listed in Table 16, as follows: 1 =
high and sharp; 2 = high and rounded; 3 = low and
sharp; 4 = low and rounded; and 5 = low and blunt.
The test result showed the expected high degree
of difference between S. kirchbergensis and C.
antiquitatis in dietary mesowear signals, i.e., the
probability of the coincidence of these two groups
was negligible (p < 0.001).
The diagram summarizing the cluster analysis
(Figure 26) reflects the distribution of the fossil rhi-
noceroses of Europe and Siberia according to
dietary preferences, against the reference data set
of 27 species of extant ungulate mammals
selected by Fortelius and Solounias (2000) on the
basis of information about their dietary preferences
(Janis, 1988). The distribution of these 27 extant
ungulates in the diagram completely coincides with
other lines of paleodietary evidence (Fortelius and
Solounias 2000; Kaiser and Solounias, 2003;
Kahlke and Kaiser, 2011). The distribution of Euro-
pean S. hundsheimensis in the diagram is congru-
ent with data from Kahlke and Kaiser (2011), and
distributions of Stephanorhinus kirchbergensis and
S. hemitoechus are congruent with data from van
Asperen and Kahlke (2014). The diagram illus-
trates the partition of the set of the animals exam-
ined into four groups. The extreme groups are
grazers and browsers. The browsers group
includes animals whose diet comprises no more
than 10% of grass. The grazers group includes ani-
mals whose diet is more than 90% of grass. Ani-
mals with a mixed diet are placed in the central part
of the diagram. The group that is closer to the graz-
ers consists of the mixed grazer feeders with pre-
vailing consumption of grass, compared to another
TABLE 17. Mesowear scores for the set of 27 typical extant species and for the fossil assemblages. (Continued on
next page.)
Species Locality Sources label
%
high
%
sharp
%
round
%
blunt N
Stephanorhinus
kirchbergensis
fossils from the
southeastern Western
Siberia
SK_SWS 100 41,6 58,4 0 12
Coelodonta antiquitatis CA_SWS 28 0 64 36 25
Stephanorhinus
hundsheimensis
fossils from the
Sussenborn
(Kahlke and
Kaiser, 2011)
SH_SUES
S
91.9 5.7 94.3 0 36
Stephanorhinus
hundsheimensis
fossils from the Voigtstedt SH_VOI 100 100 0 0 6
Stephanorhinus
hemitoechus
fossils from the
Bilzingsleben II
(Asperen and
Kahlke, 2014)
SHM_B 80 12 88 0 25
Stephanorhinus
kirchbergensis
SK_B 82.6 8.7 87 4.3 23
Stephanorhinus
kirchbergensis
fossils from the Weimar-
Ehringsdorf
SK_WE 89.5 28.9 71.1 0 76
Stephanorhinus
kirchbergensis
fossils from the Weimar-
Taubach
SK_WT 82.6 39.1 60.9 0 23
Stephanorhinus
hemitoechus
fossils from the UK MIS 7 SHM_U7 36.4 30 60 10 11
Stephanorhinus
kirchbergensis
SK_U7 83.3 16.7 66.7 16.6 6
Stephanorhinus
hemitoechus
fossils from the UK MIS
5e upland
SHM_U5u 85.7 9.5 90.5 0 21
Stephanorhinus
hemitoechus
fossils from the UK MIS
5e lowland
SHM_U5l 63.6 9.1 81.8 9.1 11
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
32
central group that is closer to the browsers, which
is mixed browser feeders.
The position of West Siberian Coelodonta
antiquitatis in the diagram lies adjacent to the typi-
cal extant grazer Ceratotherium simum. This is due
to very similar values of mesowear signals of these
two species of rhino, indicating the same high level
of adaptation to the rough abrasive food base.
Stephanorhinus kirchbergensis of West Siberia
appears in the same group with extant mammals
such as the impala (Aepyceros melampus), suma-
tran serow (Capricornis sumatraensis), wapiti (Cer-
vus elaphus canadensis), Grant’s gazelle (Gazella
granti), Thomson’s gazelle (Gazella thomsoni),
(Ovibos moschatus), common eland (Taurotragus
oryx), and bushbuck (Tragelaphus scriptus). This
group of animals belongs to mixed browser feed-
ers. It is caused by a small Euclidean distance
between the mesowear signals of animals from this
group. Also, specimens of S. kirchbergensis from
the beginning of the Late Pleistocene in the Euro-
pean areas of Weimar-Ehringsdorf and Weimar-
Alces alces extant species (Fortelius and
Solounias,
2000)
AA 100 100 0 0 30
Diceros bicornis DB 100 94.1 5.9 0 34
Dicerorhinus sumatrensis DS 100 80 20 0 5
Giraffa camelopardalis GC 94 73.7 26.3 0 61
Odocoileus hemionus OH 100 72.7 27.3 0 33
Odocoileus virginianus OV 100 88.8 11.2 0 18
Okapia johnstoni OJ 100 87.5 12.5 0 8
Rhinoceros sondaicus RS 100 100 0 0 5
Ceratotherium simum cs 0 0 72 28 26
Alcelaphus buselaphus ab 57 4.4 67.6 28 76
Bison bison bb 0 0 26.7 73.3 15
Connochaetes taurinus ct 55 15.3 55.7 29 52
Damaliscus lunatus dl 20 20 60 20 5
Equus burchelli eb 0 27 39.4 33.6 122
Equus grevyi eg 0 34.4 41.4 24.2 29
Hippotragus equinus he 85 3.9 96.1 0 26
Hippotragus niger hn 85 0 85 15 20
Kobus ellipsiprymnus ke 96 0 100 0 22
Redunca redunca rr 91 6.4 91 2.6 77
Aepyceros melampus Me 100 35.3 64.7 0 17
Capricornis sumatraensis Ca 100 45.5 50 4.5 22
Cervus elaphus canadensis Cc 100 47.4 52.6 0 19
Gazella granti Gg 88 50 50 0 18
Gazella thomsoni Gt 88 55.5 43.2 1.3 146
Ovibos moschatus Om 81 57.6 42.4 0 52
Taurotragus oryx To 100 50 50 0 14
Tragelaphus scriptus Ts 100 51 49 0 47
Species Locality Sources label
%
high
%
sharp
%
round
%
blunt N
TABLE 17 (continued).
PALAEO-ELECTRONICA.ORG
33
Taubach also belong to this group (van Asperen
and Kahlke, 2014).
BIOGEOCHEMICAL ANALYSES
To date, a great deal of information is known
about the biogeochemistry of Quaternary mammal
remains and particularly about rhinos. But research
on the Stephanorhinus genus is rare (Palmqvist et
al., 2003), and even less information is presented
specifically on S. kirchbergensis (Pushkina et al.,
2014). Comparative study of the biogeochemical
composition (including stable isotopes) of bone tis-
sue was made for the Sk_ui1 sample and a num-
ber of samples of Late and Middle Pleistocene
mammals from Middle Irtysh and other regions
because of uncertainty about the findings’ age
(probably Middle Pleistocene). We have estab-
FIGURE 26. Hierarchical cluster diagram based on the reference tooth positions of upper P4-M3 according to the
extended mesowear method (Kaiser and Solounias, 2003). Distances = Euclidean distance (root-mean-squared dif-
ference). Clusters are based on a set of 27 typical extant species model. Classification follows the conservative
(CONS) scheme of Fortelius and Solounias (2000): Browsers (CONS): AA = Alces alces, DB = Diceros bicornis, DS =
Dicerorhinus sumatrensis, GC = Giraffa camelopardalis, OH = Odocoileus hemionus, OJ = Okapia johnstoni, OV =
Odocoileus virginianus, RS = Rhinoceros sondaicus, Grazers (CONS): ab = Alcelaphus buselaphus, bb = Bison
bison, cs = Ceratotherium simum, ct = Connochaetes taurinus, dl = Damaliscus lunatus, eb = Equus burchelli, eg =
Equus grevyi, he = Hippotragus equinus, hn = Hippotragus niger, ke = Kobus ellipsiprymnus, rr = Redunca redunca;
Mixed feeders (CONS): Cc = Cervus elaphus canadensis, Ca = Capricornis sumatraensis, Gg = Gazella granti, Gt =
Gazella thomsoni, Me = Aepyceros melampus, Om = Ovibos moschatus, To = Taurotragus oryx, Ts = Tragelaphus
scriptus. European fossil populations of Stephanorhinus hundsheimensis (Kahlke and Kaiser, 2011): SH_SUESS =
Su ¨ßenborn, SH_VOI = Voigtstedt; S. kirchbergensis (van Asperen and Kahlke, 2014): SK_B = Bilzingsleben II,
SK_WE = Weimar-Ehringsdorf, SK_WT = Weimar-Taubach, SK_U7 = UK MIS 7; S. hemitoechus (van Asperen and
Kahlke, 2014): SHM_B = Bilzingsleben II, SHM_U7 = UK MIS 7, SHM_U5u = UK MIS 5e upland, SHM_U5l = UK MIS
5e lowland. Southeast Western Siberia fossil populations: SK_SWS = S. kirchbergensis, CA_SWS = Coelodonta
antiquitatis.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
34
lished that the amount of trace elements in the
Sk_ui1 tissue sample from the Ust-Ishim area (0.2
mass %) falls into the same group with suspected
or known Late Pleistocene remnants (0.1-0.25
mass %) and is significantly different from those of
the Middle Pleistocene (0.35–0.4 mass %).The
Sk_ui1 sample also displayed a quite large organic
carbon content (11.42 mass %), typical for Late
Pleistocene samples. The stable isotopic analysis
of Sk_ui1 gave the following results: 1) for bone
bioapatite, δ13СPDB = –11.30‰; and 2) for bone
collagen, δ13СPDB = –19.66‰, δ15Nair = 2.84‰.
The stable isotopic composition of the bone sam-
ple of a woolly rhinoceros from Ust-Ishim region
was: 1) for bone bioapatite, δ13СPDB = –9.40‰;
and 2) for bone collagen, δ13СPDB = –19.33‰,
δ15Nair = 5.40‰. Parameters of mammalian tis-
sues’ stable isotopic composition are determined
by isotope fractionation in the food and water,
which changes naturally through the food chains
from the producers with a different type of photo-
synthesis and preferred habitats to consumers of
higher trophic levels (Bocherens and Drucker,
2013). Interpreting the stable isotopic composition
of the tissues is more effective when there is com-
plex evidence for several species and many speci-
mens of each species, because individual values
vary, and the total ensemble of data may be shifted
in isotopic valuesdepending on the geochemical
and climatic conditions of the region (Bocherens,
2003). However, the interpretation of individual
measurements is also possible by using the accu-
mulated information. The degree of enrichment of
collagen by δ15N isotope is very informative. Low
values of δ15N in collagen of herbivores can be
explained by several factors: the use of food that is
pioneer vegetation, increased acidity of soils, and
the predominance of shrubs and trees in the diet
spectrum (Bocherens, 2003). This is in agreement
with the traditional view about the browsing feeding
strategy of S. kirchbergensis and the mesowear
analysis results for Siberian specimens. For
instance, significantly reduced δ15N relative values
are characteristic of browsers such as Alces alces,
Cervus elaphus, and Rangifer tarandus. The value
of δ15N for Sk_ui1 is smaller than that for the
woolly rhinoceros from Belgium, France, and Yaku-
tia (Bocherens, 2015), while the comparative sam-
ple of woolly rhinoceros from the Irtysh Valley
corresponds to the values for European represen-
tatives of this species. This discrepancy may reflect
different nutritional adaptation of two species of
Siberian rhinoceroses. High δ13C value (as in both
test samples) is characteristic for inhabitants of
open ecosystems where there are dominant plants
with C3 type of photosynthesis and/or an abun-
dance of lichens in the vegetation cover. The stable
isotopic composition of bioapatite of both S. kirch-
bergensis and C. antiquitatis suggests that they
consumed very fresh, probably largely melt, water.
MORPHOMETRIC ANALYSIS OF THE
JUVENILE MANDIBLE
At present, juvenile mandibles of Stephanorhi-
nus kirchbergensis are known in several localities
in Europe, China (Shennongjia) (Kahlke, 1975,
1977; Guérin, 1980; Lacombat, 2006; Tong and
Wu, 2010), and Altai (Russia) (NSMLL-12). The
largest number of juvenile mandibles is known in
the localities of Weimar-Taubach and Weimar-
Ehringsdorf in Europe. A representative collection
of Coelodonta antiquitatis juvenile mandibles of dif-
ferent individual ages from the following localities
was used as comparative material: Krasniy Yar of
Tomsk Province (Shpansky, 2014), Novosibirsk
Province, and Altai Territory.
The main differences between juvenile mandi-
bles of S. kirchbergensis and those of Coelodonta
antiquitatis are the absence of thickening in the
ventral part of the horizontal corpus and more mas-
sive teeth (Table 2).
The change in the average indices of lower
deciduous teeth (relation of tooth width to its
length) from dp1 to dp4 is not linear in Stephanorhi-
nus kirchbergensis or in Coelodonta antiquitatis
(Figure 27). At the same time, the main trend in C.
antiquitatis from West Siberia is that the greatest
width index is observed on dp2, while dp4 has less
width index than dp2, and it has often the minimal
value of width index with respect to the other teeth
of the same jaw (Figure 28). Dynamics of changes
in the average values of the width indices of the
deciduous teeth from dp1 to dp4 in C. antiquitatis
from West Siberia, as well as from European
regions, reflects this general trend. The exception
is the value for dp1 in the European plot, which
was obtained from the only specimen, and there-
fore may not reflect the value of the population
(Figure 27). For the mandible of S. kirchbergensis
(NSMLL-12), another index distribution is
observed. The width index of dp4 is the highest. It
is much greater than the width index of dp2 (Figure
29). The same may be observed for four mandibles
from Weimar-Taubach (MIS 5e) (Kahlke, 1977),
where the width index of all dp4 is at least the
same as that of dp2 (Figure 29). For the older
locality of Weimar-Ehringsdorf (MIS 7) (Kahlke,
PALAEO-ELECTRONICA.ORG
35
1975), the width index of dp4 compared to dp2 is
more variable. The maximal value of the width
index is observed on dp1, but the main trend in the
width index of dp4 and dp2 remains the same, i.e.,
for three of five jaws that we present here, the
width index of dp4 is lower than that of dp2. In the
other two jaws, the width index between dp4 and
dp2 is close to that in C. antiquitatis (Figure 29).
Considering that mesowear signals of adults from
the locality of Weimar-Ehringsdorf (MIS 7) on the
hierarchical graph (Figure 26) also shift towards
grazers, the slight shift of teeth proportions in juve-
niles towards the C. antiquitatis condition appears
quite explainable. The trend of changes in decidu-
ous teeth indices from dp1 to dp4 in NSMLL-12 is
similar with the trend of change in the indices of S.
kirchbergensis average indices from Europe (Fig-
ure 27). For localities in China, we can only
observe the general dynamics of the average indi-
ces of the deciduous teeth from dp1 to dp4 for S.
kirchbergensis (Figure 27), because Tong and Wu
(2010) provided information only on isolated teeth
and there is no full information on mandibles of
juveniles.
To determine the age of the juvenile NSMLL-
12, a juvenile individuals of extant African black rhi-
nos (Diceros bicornis) and white rhinos (Ceratothe-
rium simum) were used as comparative material
(Schaurte, 1966; Dittrich, 1974), and the method of
age determination proposed by Hillman-Smith et
al. (1986) was used. Based on the assumption that
the order of deciduous teeth eruption, functioning,
and replacement were close to those of extant rhi-
noceroses, we applied the method introduced by
Hillman-Smith et al. (1986) in order to estimate the
age of NSMLL-12. In NSMLL-12, only the decidu-
ous teeth dp2 (STL5), dp3 (STL5), and dp4 (STL4)
were erupted and functioned in the initial stage.
The degree of wear of preserved deciduous teeth
of NSMLL-12 is closest to the specimen IQW 1968/
9761 of the juvenile mandible of S. kirchbergensis
from Taubach (Kahlke, 1977). Therefore, dp1 in
NSMLL-12 was not used, or dp1 was used in the
earliest stage (STL2 or STL3), just as in IQW 1968/
9761. Because alveolar pockets for permanent
teeth P2 and P3 were not found after apertum of
the NSMLL-12 textus, it is likely that m1 had not yet
started erupting from the bone tissue. It corre-
sponds to stages STL1 or STL2. Information about
stages of wear of dp1-m1 from NSMLL-12 com-
pletely corresponds to information on the expected
state of mandibular teeth in Hillman-Smith et al.
(1986: table IV). This corresponds to the age of a
juvenile rhinoceros, 12–18 months.
DISCUSSION
Sizes and morphological features of lower
dentitions and mandibles from different locations in
Siberia correspond to specimens known from
Europe. The adult mandibles studied may be sepa-
rated into two groups, which differ considerably by
the length of their tooth rows. The first group,
including the specimens IAE KY-4323, IRM 2436,
and GR PC 1165, is characterized by the length of
р2-m3 varying from 263 mm to 265 mm. The sec-
ond group of specimens, NSMLL-10, Sk_ui1,
NSMLL 22090, GR PC 1164, and KB MAN K-397,
has a length of tooth row that varies from 278 mm
FIGURE 27. Indexes of the lower deciduous teeth of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from
Siberian, European, and Chinese regions.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
36
to 289 mm. Premolar tooth row р2-4 length for all
mandibles varies from 108 mm to 118 mm (Table
4). Differences are more visible in the length of m1-
3. For the first group, the length of m1-3 varies
from 152 mm to 155 mm, while for the second it
varies from 163 mm to 171 mm. It may be
assumed that these differences are a consequence
of expressed sexual dimorphism of body size for
Stephanorhinus kirchbergensis, by analogy with
sexual dimorphism of body size in extant species:
Rhinoceros unicornis (Dinerstein, 1991) and Cera-
totherium simum (Owen-Smith, 1975). Thus, of the
total number of jaws considered, 30% are jaws with
short dentition (presumably female), 20% are
young individuals (less than 10 years old), and
50% are jaws with a long dentition (presumably
male). All jaws with a short tooth row belong to
adult animals older than 20 years, while there are
two groups for the jaws with a long dentition: young
animals aged 15–20 years and individuals over 20
years old. To a certain extent this distribution
agrees with the general model of sexual dimor-
phism of extant rhinos that reflects the hierarchy of
dominance (Mihlbachler, 2003, 2005, 2007). This
model explains the resurgence in mortality among
young rhino males because of the severe domi-
nance of older individuals. Such a surge in mortal-
ity is not observed among females. A more detailed
FIGURE 28. Indexes of the lower deciduous teeth dp1-dp4 of Coelodonta antiquitatis (Ca).
PALAEO-ELECTRONICA.ORG
37
study of the model of sexual dimorphism of S.
kirchbergensis using the capabilities of statistical
systems is currently impossible because of the
insufficient amount of material. For the same rea-
son, it has been impossible to study the morphol-
ogy of sexual dimorphism, because for definitive
studies it is necessary to have complete skeletons
of S. kirchbergensis individuals of both sexes, or at
least complete skulls with mandibles, which to date
have not been found.
Comparative analysis of the postcranium from
the localities of West Siberia showed their morpho-
logical similarity and closeness in size to remains
of Stephanorhinus kirchbergensis from Europe.
Morphological analysis of the dental features
presented and comparison with extant typical
browsers and grazers (Fortelius, 1982; Clauss et
al., 2008) show the shift towards the browsers in
the material studied. On the other hand, it would be
an oversimplification to assume that Stephanorhi-
nus kirchbergensis was exclusively a forest
dweller. The diet of S. kirchbergensis contained a
high percentage of abrasive food, as is observed
from mesowear analysis of European remains
(Hernesniemi et al., 2011; Kahlke and Kaiser, 2011;
FIGURE 29. Indexes of the lower deciduous teeth dp1-dp4 of Stephanorhinus kirchbergensis (Sk).
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
38
van Asperen and Kahlke, 2014); these analyses
showed significant dietary flexibility that depended
on the quality of the ecological environment. At the
same time, morphological features and mesowear
signals of S. kirchbergensis inhabiting Europe in
the Late Pleistocene indicate a higher dietary spe-
cialization shifted to browsers, compared with the
early and Middle Pleistocene. Mesowear analysis
of the dietary preferences of Siberian S. kirchber-
gensis also shows their higher dietary specializa-
tion, comparable to mesowear signals of S.
kirchbergensis from the beginning of the Late
Pleistocene in Europe from Weimar-Taubach (van
Asperen and Kahlke, 2014). However, the heavy
continental climate of Siberian regions and the
poorer food base contributed to the process of S.
kirchbergensis adaptation to the local environment.
In addition to the increased molar row length, char-
acteristic of S. kirchbergensis in all regions of their
habitat, representatives of the Siberian region tend
to increase the relative width of the upper and
lower molars (Figures 30, 31). Thus, the increase
in the occlusal surface area is observed. It allowed
increasing the volume of the processed food per
unit of time and adapting to the poor food base.
According to the results of mesowear analysis,
unlike Coelodonta antiquitatis, S. kirchbergensis
could not inhabit West Siberia in the periods of tun-
dra–steppe habitat conditions. The structure and
relief of the occlusal surface of teeth show that S.
kirchbergensis had not adapted to using strongly
abrasive vegetation. Even in the interglacial peri-
ods, it probably had to disperse to the more south-
erly areas, for the same reasons. The results of the
stable isotopic study of the single sample of the
Siberian S. kirchbergensis indicate that it was a
browser-type that fed presumably in predominantly
open landscapes.
Despite overall similar mesowear signals for
Stephanorhinus kirchbergensis in the Siberian
region and in Europe at the beginning of Late
Pleistocene (MIS 5e), it cannot be stated with cer-
tainty that S. kirchbergensis existed contempora-
neously in Siberia and Europe. A variety of
landscape and climatic conditions and interspecies
competition had led to significant variability of
Pleistocene rhinoceroses’ feeding behavior (van
Asperen and Kahlke, 2014). This circumstance
does not allow extrapolating the paleoecological
assessments gained from the European materials
to Siberian representatives of S. kirchbergensis.
On one hand, all the remains of S. kirchbergensis
previously found in southeastern West Siberia
(Kemerovo and Tomsk Provinces) (Alekseeva,
1980; Shpansky and Billia, 2012) and on Vilyuy
River (Dubrovo, 1957) were assigned to the Middle
Pleistocene, based on accompanying fauna,
including Mammuthus ex gr. trogontherii-chosari-
cus, Megaloceros giganteus ruffi, and Equus ex gr.
mosbachensis-germanicus. In addition, it can be
noted that the degree of fossilization of the existing
Middle Pleistocene residues agrees to a certain
extent with the degree of fossilization of new
remains of S. kirchbergensis from the Altai Territory
and Novosibirsk Province. On the other hand,
although the latest findings of S. kirchbergensis on
the beaches from Altay Territory and Novosibirsk
Province do not have a stratigraphic linkage, it is
FIGURE 30. Indexes of the upper permanent teeth of Stephanorhinus kirchbergensis and Coelodonta antiquitatis
from Siberian, European, and Chinese regions.
PALAEO-ELECTRONICA.ORG
39
characteristic that the accompanying fauna of all
these locations is mainly from the Late Pleisto-
cene, and there are very few remains of the Middle
Pleistocene megafauna.
However, until now, none of the finds related
to Stephanorhinus kirchbergensis discovered in
Siberia has been unequivocally attributed to the
Late Pleistocene. Quantitative data on the chemi-
cal composition of the sample from the Middle
Irtysh give an indirect indication that individual
specimens of S. kirchbergensis could have existed
in West Siberia in the Late Pleistocene. To date,
we can assert that despite its small number, the
population of S. kirchbergensis was distributed
fairly widely in Siberia in the places most favorable
to their dietary preferences—river valleys with rich
bush vegetation.
ACKNOWLEDGMENTS
The authors are thankful to E.M. Matskevich,
the head of the Museum of Arts of the North Peo-
ples (Kargasok settlement, Tomsk Province); I.V.
Orlova, main curator of the Novosibirsk State
Museum of Local History; V.K. Kreshik, assistant to
the chief curator of Novosibirsk State Museum of
Local History; I.N. Shitikova, head of the Nature
Department of the Irkutsk Local History Museum;
N.V. Peristov, head of the studio Arhaika; and A.L.
Dorogov for access to collections of Stephanorhi-
nus kirchbergensis remains. This study (research
grants No 8.1.80.2015) was supported by the
Tomsk State University Academic D. I. Mendeleev
Fund Program in 2015. The authors thank S.
Ivantsov, Tomsk State University, for translation,
and J. Kollantai, Tomsk State University, for style
review.
REFERENCES
Alekseeva, E.V. 1980. Mlekopitayushchie Pleistocena yugo-vostoka Zapadnoi Sibiri. Nauka,
Moskow. [In Russian]
Belyaeva, E.I. 1940. Novye nakhodki ostatkov nosoroga Mercka na territorii SSSR. Priroda,
8:82. [In Russian]
Belyaeva, E.I. 1966. Semeistvo Rhinocerotidae Owen, 1845. In Mlekopitayushchie
eopleistocena Zapadnogo Zabaikalia. Trudy Geologicheskogo instituta Akademii Nauk
SSSR, 152:92-143. [In Russian]
Billia, E.M.E. 2007. First records of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia,
Rhinocerotidae) from the Kuznetsk Basin (Kemerovo region, Kuzbass area, South-East of
Western Siberia). Bollettino della Società Paleontologica Italiana, 46:95-100.
FIGURE 31. Indexes of the lower permanent teeth of Stephanorhinus kirchbergensis and Coelodonta antiquitatis from
Siberian, European, and Chinese regions.
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
40
Billia, E.M.E. 2008a. The skull of Stephanorhinus kirchbergensis (Jäger 1839) (Mammalia,
Rhinocerotidae) from the Irkutsk region (Southwest Eastern Siberia). Quaternary
International, 179:20-24. https://doi.org/10.1016/j.quaint.2007.08.034
Billia, E.M.E. 2008b. Revision of the fossil material attributed to Stephanorhinus kirchbergensis
(Jäger 1839) (Mammalia, Rhinocerotidae) preserved in the museum collections of the
Russian Federation. Quaternary International, 179:25-37. https://doi.org/10.1016/
j.quaint.2007.09.034
Billia, E.M.E. 2011a. Siti paleontologici a “Rinoceronte di Merck”, Stephanorhinus kirchbergensis
(Jäger, 1839) (Mammalia, Perissodactyla), in Istria, Quarnero e Dalmazia [Paleontoloska
Najdisca Merckovega Nosoroga, S. kirchbergensis (Jäger, 1839) (Mammalia,
Perissodactyla) v Istri in Dalmaciji – Paleontoloska nalazista Merckovog Nosoroga, S.
kirchbergensis (Jäger, 1839) (Mammalia, Perissodactyla), u Istri i Dalmaciji]. Atti - Centro di
Ricerche Storiche / Centar za Povijesna Istrazivanja / Sredisce za Zgodovinska
Raziskovanja, Rovigno/Rovinij, Croatia/Hrvatska, XLI:9-31. (with English, Croatian &
Slovenian abstracts)
Billia, E.M.E. 2011b. Occurrences of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia,
Rhinocerotidae) in Eurasia – An account. Acta Palaeontologica Romaniae, 7:17-40.
Bocherens, H. 2003. Isotopic biogeochemistry and the paleoecology of the mammoth steppe
fauna. DEINSEA, 9:57-76.
Bocherens, H. 2015. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe.
Quaternary Science Reviews, 117:42-71. https://doi.org/10.1016/j.quascirev.2015.03.018
Bocherens, H. and Drucker, D.G. 2013. Terrestrial teeth and bones, p. 304-314. In Elias, S.A.
and Mock, C.J. (eds.), Encyclopedia of Quaternary Science. 2nd edition. Elsevier, Edinburgh.
https://doi.org/10.1016/b978-0-444-53643-3.00341-1
Chersky, I.D. 1874. Opisanie cherepa nosoroga, razlichnago ot’ Rhinoceros tichorhinus. Zapiski
Imperatorskoy Akademii Nauk, 25:65-75. [In Russian]
Chow, B.-S. 1979. The fossil rhinocerotides of locality 1, Choukoutien. Vertebrata PalAsiatica,
17:236-258. [In Chinese with English summary]
Clauss, M., Kaiser, T., and Hummel, J. 2008. The morphophysiological adaptations of browsing
and grazing mammals, p. 47-88. In Gordon, I.J. and Prins, H.H.T. (eds.), The Ecology of
Browsing and Grazing. Springer, Berlin. https://doi.org/10.1007/978-3-540-72422-3_3
David, I.A., Tatarinov, K.A., and Svistun, V.I. 1990. Khishchnye, khobotnye i kopytnye rannego
Pleistocena yugo-zapada SSSR. Shtiinca, Kishinyov. [In Russian]
Dinerstein, E. 1991. Sexual dimorphism in the greater one-horned rhinoceros (Rhinoceros
unicornis). Journal of Mammalogy, 72:450-457. https://doi.org/10.2307/1382127
Dittrich, L. 1974. Beobachtungen zum Milchzahndurchbruch beim Spitzmaul-(Diceros bicornis)
und Breitmaulnashorn (Ceratotherium simum). Säugetierkundliche Mitteilungen, 22:289-295.
Dubrovo, I.A. 1957. Ob ostatkakh Parelephas wüsti (Pawl.) i Rhinoceros mercki Jaeger iz
Yakutii. Bjulletin Komissii Izucheniya Chetvertichnogo Perioda, 21:97-104. [In Russian]
Fortelius, M. 1982. Ecological aspects of dental functional morphology in the Plio-Pleistocene
rhinoceroses of Europe, p. 163-181. In Kurtén, B. (ed.), Teeth: Form, Function, and
Evolution. Columbia University Press, New York.
Fortelius, M., Mazza, P., and Sala, B. 1993. Stephanorhinus (Mammalia: Rhinocerotidae) of the
western European Pleistocene, with a revision of S. etruscus (Falconer, 1868).
Palaeontographia Italica, 80:63-155.
Fortelius, M. and Solounias, N. 2000. Functional characterization of ungulate molars using the
abrasion-attrition wear gradient: a new method for reconstructing paleodiets. American
Museum Novitates, 3301:1-36. https://doi.org/10.1206/0003-
0082(2000)301%3C0001:fcoumu%3E2.0.co;2
Gromova, V.I. 1935. Ob ostatkakh nosoroga Merka (Rhinoceros mercki Jaeg.) s nizhnei Volgi.
Trudy Paleozoologicheskogo Instituta Akademii Nauk SSSR, 4:91-136. [In Russian]
Gromova, V.I. 1950. Opredelitel mlekopitayushchikh SSSR po kostyam skeleta. Trudy komissii
po izucheniyu chetvertichnogo perioda, 9(1):1-240. [In Russian]
Guérin, C. 1980. Les rhinocéros (Mammalia, Perissodactyla) du Miocène terminal au
Pléistocène supérieur en Europe occidentale: comparaison avec les espèces actuelles.
Documents des Laboratoires de Géologie Lyon, 79:1-421.
Hernesniemi, E., Blomstedt, K., and Fortelius, M. 2011. Multi-view stereo three-dimensional
reconstruction of lower molars of Recent and Pleistocene rhinoceroses for mesowear
analysis. Palaeontologia Electronica, 14.2.2T.
PALAEO-ELECTRONICA.ORG
41
Hillman-Smith, A.K.K., Owen-Smith, N., Anderson, J.L., Hall-Martin, A.J., and Selaladi, J.P.
1986. Age estimation of the white rhinoceros (Ceratotherium simum). Journal of Zoology,
21 0:355 -379 . htt ps:/ /doi .org/ 10.1111/ j.14 69-79 98.19 86.t b03639.x
Hitchins, P.M. 1978. Age determination of the black rhinoceros (Diceros bicornis Linn.) in
Zululand. South African Journal of Wildlife Research, 8:71-80.
Jäger, G.F. 1835–39. Über die fossilen Säugetiere welche in Würtemberg in verschiedenen
Formationen aufgefunden worden sind, nebst geognotischen Bemerkungen über diese
Formationen. C. Erhard Verlag, Stuttgart.
Janis, C.M. 1988. An estimation of tooth volume and hypsodonty indices in ungulate mammals,
and the correlation of these factors with dietary preferences. Mémoires du Muséum National
d’Histoire Naturelle, 53:367-387.
Kahlke, H.-D. 1975. Die Rhinocerotiden-Reste aus den Travertinen von Weimar-Ehringsdorf.
Palaontologische Abhandlungen, 23:337-397.
Kahlke, H-D. 1977. Die Rhinocerotidenreste aus den Travertinen von Taubach.
Quartärpaläontologie, 2:305-359.
Kahlke, R.-D. and Kaiser, T.M. 2011. Generalism as a subsistence strategy: advantages and
limitations of the highly flexible feeding traits of Pleistocene Stephanorhinus hundsheimensis
(Rhinocerotidae, Mammalia). Quaternary Science Reviews, 30:2250-2261. https://doi.org/
10.1016/j.quascirev.2009.12.012
Kaiser, T.M. and Solounias, N. 2003. Extending the tooth mesowear method to extinct and extant
equids. Geodiversitas, 25:321-345.
Krivonogov S.K. 1988. [Stratigraphy and paleogeography of lower Irtysh valley in the last
glaciation (on carpological data)]. Nauka, Novosibirsk. [In Russian]
Lacombat, F. 2005. Les rhinocéros fossiles des sites préhistoriques de l'Europe
méditerranéenne et du Massif Central. Paléontologie et implications biochronologiques. BAR
International Series, 1419:1-185.
Lacombat, F. 2006. Morphological and biometrical differentiation of the teeth from Pleistocene
species of Stephanorhinus (Mammalia, Perissodactyla, Rhinocerotidae) in Mediterranean
Europe and the Massif Central, France. Palaeontographica Abteilung A, 274:71-111.
Lobachev, Y.V., Vasiliev, S.K., and Orlova, L.A. 2012. Pozdnepleistocenovaya teriofauna s reki
Chumysh (Altaisky krai) i novye dannye po mestonakhozdeniyu na reke Chik
(Novosibirskaya oblast). Problemy arkheologii, etnografii, antropologii Sibiri i sopredelnykh
territoryi, 18:106-110. [In Russian]
Lobachev, Y.V., Lobachev, A.Y., and Billia, E.M.E. 2014. Morfologicheskie i biomekhanicheskie
osobennosti zhevatelnogo apparata Coelodonta antiquitatis (Blumenbach, 1799) i
Stephanorhinus kirchbergensis (Jäger, 1839), p. 168-170. In Diversifikaciya i etapnost
evolyucii organicheskogo mira v svete paleontologicheskoi letopisi. Sankt-Peterburg. [In
Russian]
Martynov, V.A., Mizerov, B.V., Nikitin, V.P., and Shaevich, Y.E. 1977. Geomorphologicheskie
structury reki Ob okrestnostei Novosibirska. [In Russian]
Mihlbachler, M.C. 2003. Demography of late Miocene rhinoceroses (Teleoceras proterum and
Aphelops malacorhinus) from Florida: linking mortality and sociality in fossil assemblages.
Paleobiology, 29:412-428. https://doi.org/10.1666/0094-
8373(2003)029%3C0412:dolmrt%3E2.0.co;2
Mihlbachler, M.C. 2005. Linking sexual dimorphism and sociality in rhinoceroses: insights from
Teleoceras proterum and Aphelops malacorhinus from the late Miocene of Florida. Bulletin of
the Florida Museum of Natural History, 45:495-520.
Mihlbachler, M.C. 2007. Sexual dimorphism and mortality bias in a small Miocene North
American rhino, Menoceras arikarense: insights into the coevolution of sexual dimorphism
and sociality in rhinos. Journal of Mammalian Evolution, 14:217-238. https://doi.org/10.1007/
s10914-007-9048-4
Owen-Smith, R.N. 1975. The social ethology of the white rhinoceros Ceratotherium simum
(Burchell, 1817). Zeitschrift für Tierpsychologie, 38:337-384. https://doi.org/10.1111/j.1439-
0310.1975.tb02010.x
Palmqvist, P., Gröcke, D.R., Arribas, A., and Fariña, R.A. 2003. Paleoecological reconstruction
of a lower Pleistocene large mammal community using biogeochemical (δ13C, δ15N, δ18O,
Sr:Zn) and ecomorphological approaches. Paleobiology, 29:205-229. https://doi.org/10.1017/
s0094837300018078
LOBACHEV, ET AL.: STEPHANORHINUS KIRCHBERGENSIS IN SIBERIA
42
Panychev, V.A. 1979. Radiouglerodnaya chronologiya alluvialnykh otlozhenyi Predaltaiskoi
ravniny. Nauka, Novosibirsk. [In Russian]
Popowics, T.E. and Fortelius, M. 1997. On the cutting edge: tooth blade sharpness in
herbivorous and faunivorous mammals. Annales Zoologici Fennici, 34:73-88.
Pushkina, D., Bocherens, H., and Ziegler, R. 2014. Unexpected palaeoecological features of the
Middle and Late Pleistocene large herbivores in southwestern Germany revealed by stable
isotopic abundances in tooth enamel. Quaternary International, 339-340:164-178. https://
doi.org/10.1016/j.quaint.2013.12.033
Rusanov, G.G. and Orlova, L.A. 2013. Radiouglerodnye datirovki (SBRAS) Gornogo Altaya i
Predaltayskoi ravniny. Biysk. [In Russian]
Sanson, G. 2006. The biomechanics of browsing and grazing. American Journal of Botany,
93:1531-1545. https://doi.org/10.3732/ajb.93.10.1531
Schaurte, W. 1966. Beiträge zur Kenntnis des Gebisses und Zahnbaues der afrikanischen
Nashörner. Säugetierkundliche Mitteilungen, 14:327-341.
Shpansky, A.V. and Billia, E.M.E. 2012. Records of Stephanorhinus kirchbergensis (Jäger, 1839)
(Mammalia, Rhinocerotidae) from the Ob’ River at Krasny Yar (Tomsk region, southeast of
Western Siberia). Russian Journal of Theriology, 11:47-55.
Shpansky, A.V. 2014. Juvenile remains of the “woolly rhinoceros” Coelodonta antiquitatis
(Blumenbach 1799) (Mammalia, Rhinocerotidae) from the Tomsk Priob’e area (southeast
Western Siberia). Quaternary International, 333:86-99. https://doi.org/10.1016/
j.quaint.2014.01.047
Shpansky A.V. 2016. New finds of Merck Rhinoceros (Stephanorhinus kirchbergensis Jäger
1839) (Rhinocerotidae, Mammalia) in Ob Area, Tomsk region. Geosphere Research, 1:24-
39. [In Russian] https://doi.org/10.17223/25421379/1/3
Shpansky, A.V., Svyatko, S.V., Reimer, P.J., and Titov, S.V. 2016. Finds of skeletons of Bison
priscus Bojanus (Artiodactyla, Bovidae) from Western Siberia. Russian Journal of Theriology,
15:100-120. https://doi.org/10.15298/rusjtheriol.15.2.04
Silaev, V.I., Ponomarev, D.V., Slepchenko, S.M., Bondarev, A.A., Kiselеva, D.V, Smoleva, I.V.,
and Khazov, A.F. 2015. Mineralogical and geochemical studies of bone detritus of
Pleistocene mammals, including the earliest in Northern Eurasia Humans. Bulletin of Perm
University. Geology, 4:6-31. https://doi.org/10.17072/psu.geol.29.6
Steuer, P., Clauss, M., Südekum, K.-H., Hatt, J.-M., Silinski, S., Klomburg, S., Zimmermann, W.,
Fickel, J., Streich, W.J., and Hummel, J. 2010. Comparative investigations on digestion in
grazing (Ceratotherium simum) and browsing (Diceros bicornis) rhinoceroses. Comparative
Biochemistry and Physiology. Part A: Molecular & Integrative Physiology, 156:380-388.
https://doi.org/10.1016/j.cbpa.2010.03.006
Tong, H.W. and Wu, X.Z. 2010. Stephanorhinus kirchbergensis (Rhinocerotidae, Mammalia)
from the Rhino Cave in Shennongjia, Hubei. Chinese Science Bulletin, 55:1157-1168. https://
doi.org/10.1007/s11434-010-0050-5
van Asperen, E.N. and Kahlke, R.-D. 2014. Dietary variation and overlap in Central and
Northwest European Stephanorhinus kirchbergensis and S. hemitoechus (Rhinocerotidae,
Mammalia) influenced by habitat diversity. Quaternary Science Reviews, 107:47-61. https://
doi.org/10.1016/j.quascirev.2014.10.001
Vasiliev, S.K. and Orlova, L.A. 2006. K voprosu o Taradanovskogo mestonakhozhdeniya fauny
krupnykh mlekopitayushch. Problemy arkheologii, etnografii, antropologii Sibiri i
sopredelnykh territoryi, 12(1):36-42. [In Russian]
Vasiliev, S.K., Lobachev, Y.V., and Lobachev, A.Y. 2014. Novye dannye po mestonakhozdeniyam
pozdnepleistocenovoi megafauny na rekakh Chumysh i Chick (Altaiskyi krai i Novosibirskaya
oblast). Problemy arkheologii, etnografii, antropologii Sibiri i sopredelnykh territoryi, 15:15-
18. [In Russian]
Volkov, I.A. and Arhipov, S.A. 1978. Chetvertichnye otlozheniya raiona Novosibirska. Izdatelstvo
IGiG, Novosibirsk. [In Russian]
Volkova, V.S. and Babushkin, A.E. (eds.) 2000. Uniphicirovannaya stratigraficheskaya schema
chetvertichnykh otlozheni Zapadno-Sibirskoi ravniny. SNIIGMS, Novosibirsk. [In Russian]
