GeolCarp_Vol67_No1_69

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, FEBRUARY 2016, 67, 1, 69—82 doi: 10.1515/geoca-2016-0004

Rhinocerotidae from the Upper Miocene deposits

of the Western Pannonian Basin (Hungary):

implications for migration routes and biogeography

LUCA PANDOLFI

, MIHÁLY GASPARIK

and IMRE MAGYAR

3,4

University of Roma Tre, Department of Sciences, section of Geology, Largo S.L. Murialdo 1, 00186 Rome, Italy;

luca.pandolfi@uniroma3.it

Hungarian Natural History Museum, Department of Palaeontology and Geology, H-1431 Budapest, Pf. 137, Hungary;

gasparik@nhmus.hu

MTA-MTM-ELTE Research Group for Paleontology, H-1431 Budapest, Pf. 137, Hungary;

MOL Hungarian Oil and Gas Plc., H-1117 Budapest, Október 23. u. 18, Hungary; immagyar@mol.hu

(Manuscript received April 22, 2015; accepted in revised form December 8, 2015)

Abstract:

Although the rhinoceros remains have high biochronological significance, they are poorly known or scarcely

documented in the uppermost Miocene deposits of Europe. Several specimens collected from the Upper Miocene (around
7.0 Ma, Turolian) deposits of Kávás (Pannonian Basin, Western Hungary), previously determined as Rhinoceros sp.,
are revised and described in this paper. The postcranial remains of these specimens belong to “Dihoplus” megarhinus
(de Christol) on the basis of the morphological and morphometric characters of humerus, radii, metacarpal and metatar-
sal elements. An overview of rhinoceros remains from several uppermost Miocene localities and the revision of the
rhinoceros material from the Pannonian Basin suggest that “D.” megarhinus spread during the latest Miocene from the
Pannonian Basin towards Italy. The occurrences of this species in Western Hungary and Italy during the latest Miocene
further imply that Rhinocerotini species were biogeographically segregated between Western, Southern and Central
Europe.

Key words: “Dihoplus” megarhinus, postcranium, paleobiogeography, biochronology, latest Miocene, Kávás, Pannon

ian

Basin.

Introduction

The  occurrences  of  Rhinocerotidae  species  have  been  fre-
quently  used  as  a  biochronological  tool  since  the  works  of
Guérin (1980, 1982). However, the temporal and spatial dis-
tribution  of  some  species  is  still  debated  or  remains  poorly
known,  as  does  their  taxonomic  status  and  morphological
variability (Guérin 1980, 2004; Groves 1983; Cerdeño 1992,
1995,  1998;  Heissig  1999;  Pandolfi  &  Tagliacozzo  2015).
During the

Late

Miocene, only three species belonging to the

tribe Rhinocerotini (sensu Heissig 1999=Rhinocerotina in
Antoine 2002) have been identified in Europe: Dihoplus
schleiermacheri (Kaup 1832), Dihoplus pikermiensis (Toula
1906) and Ceratotherium neumayri (Osborn 1900).

D. schleiermacheri occurs in the Vallesian and lower

Turolian deposits (from MN 9 to MN 12) at several Central
and Western European localities (Kaup 1832; Guérin 1980;
Cerdeño 1992; Heissig 1999). D. pikermiensis occurs in the
Turolian deposits (?MN 11—MN 13) of the Balkan Peninsula,
in particular in Greece and Bulgaria (Geraads 1988; Heissig
1999;  Geraads  &  Spassov  2009).  C.  neumayri  has  been
reported  from  several  fossiliferous  localities  (MN 10—
MN 13)  of  the  Balkan  Peninsula,  Caucasus,  Anatolia  and
Iran (Osborn 1900; Geraads 1988; Heissig 1999; Geraads &
Spassov 2009; Giaourtsakis 2009; Pandolfi 2015a). The co-
existence of  C. neumayri  and D. pikermiensis is well-docu-
mented  at  Pikermi  and  Samos  and  the  two  species  are

constantly present in Greece until the MN 12—MN 13 transi-
tion (Heissig 1996). The rhinoceros remains chronologically
referred to MN 13 are scarcely documented in Europe (Heis-
sig 1996).

During the Pliocene (MN 14, MN 15, MN 16a) the three

aforementioned species were not reported from Europe. The
tribe  Rhinocerotini  is  instead  represented  by  four  species:
“Dihoplus”  megarhinus  (de  Christol  1834),  “Stephanorhi-
nus”  miguelcrusafonti  (Guérin  &  Santafé-Llopis  1978:
which is here provisionally retained within the genus Stepha-
norhinus,  although  cranial  remains  of  this  species  are  un-
known  and  its  systematic  position  appears  questionable),
Stephanorhinus jeanvireti (Guérin 1972: following the ICZN
art. 23.12 and 23b, Rhinoceros elatus Croizet & Jobert 1828
is  synonymous  with  this  species,  details  are  reported  in
Guérin & Tsoukala 2013, p. 454) and Stephanorhinus etruscus
(Falconer 1868).

“D.” megarhinus has been considered a typical Pliocene

species (Guérin 1980; Pandolfi 2013) and it has been also re-
corded in Turkey and Russia (Guérin & Sen 1998; Fukuchi
et  al.  2009).  “S.”  miguelcrusafonti  has  a  restricted  geogra-
phical  and  chronological  range;  it  has  been  recovered  at
a few early Pliocene Spanish and French localities (Guérin &
Santafé-Llopis  1978;  Guérin  1980).  S.  jeanvireti  has  been
frequently  documented  from  Late  Pliocene  localities  of
France and Italy (Guérin 1972, 1980; Pandolfi 2013), but it
has also been recorded in Slovakia (Ďurišová 2004; Vlačiky

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et al. 2008; Šujan et al. 2013), in Romania (Guérin 1980), in
Russia (Titov 2008), and recently in Greece (Guérin &
Tsoukala 2013). At the end of the Pliocene, S. etruscus has
been also recorded in Western Europe (Guérin 1980;
Cerdeño 1993; Mazo 1995; Pandolfi 2013; Pandolfi & Marra
2015; Pandolfi et al. 2015a).

The aim o

f this paper is to describe postcranial rhinoceros

remains  collected  in  the  Upper  Miocene  deposits  at  Kávás
(Western Hungary, Fig. 1) and (1) to analyse morphological
differences  between  the  Late  Miocene  and  Pliocene  species
usually assigned to the genera Dihoplus and Stephanorhinus
and (2) to discuss the implications of these findings for mi-
gration  patterns  and  biogeography  of  Rhinocerotidae  at  the
end of the Miocene.

Geological and stratigraphic background

The specimens analysed here were collected from grey,

clayey,  fine-grained  sand  or  sandstone  deposits  near  the
village of Kávás in 1979 (although the name of the collector
is missing from the Inventory Book, it was probably Dénes
Jánossy). Kávás is located in the western part of the Neogene
Pannonian  Basin  (Fig. 1).  The  basement  of  the  Neogene
infill  in  this  region  is  represented  by  Upper  Triassic  dolo-
mites belonging to the Transdanubian Range (Bakony Mts.).
The pre-Neogene basement forms a flat platform at a depth
of ca. 1800 m below sea level, separating the Kisalföld sub-
basin to the north and the Zala subbasin to the south (Haas et
al. 2010).

The Neogene basin fill at Kávás was penetrated by a hydro-

carbon  exploration  well  (Nf-3)  in  the  early  1970s.  Its
1853 m-thick  Neogene  sequence  started  with  a  78 m-thick
marine  unit,  consisting  of  glauconitic  calcareous  marl  with
abundant remains of benthic and planktonic foraminifers and
pectinid bivalves. The fossils indicate Middle Miocene (Bade-
nian) age. The overlying unit, from 1775 m up to the surface,
belongs  to  the  Pannonian.  The  Pannonian  Stage,  as  used  in

Hungary  (Pannonian  sensu  lato),  corresponds  to  the  Upper
Miocene  and  the  Pliocene.  Its  sedimentary  succession  was
deposited in Lake Pannon, a giant brackish lake, and in the
adjacent  deltaic  and  fluvial  environments.  The  Pannonian
succession in the Nf-3 borehole can be subdivided into five
lithological  units,  including:  (1)  marls  deposited  in  a  deep
lacustrine  setting  (Endrőd  Formation;  1775  to  1623 m),
(2)  alternation  of  argillaceous  marl  and  fine-grained  sand-
stone layers, the latter deposited by turbidity currents (Szol-
nok  Formation;  1623  to  1410 m),  (3)  argillaceous  marl  and
silt  with  subordinate  sandstone  layers,  deposited  on  the
shelf-margin  slope  (Algyő  Formation;  1410  to  1160 m),
(4)  alternation  of  argillaceous  marl,  sandstone,  and  lignite
layers  deposited  in  shallow  lacustrine,  deltaic,  and  paludal
environments  (Újfalu  Formation;  1160  to  ca.  400 m),  and
(5)  clay,  sand,  and  fine-grained  gravel  deposited  in  flood
plains,  point  bars,  and  river  channels  (Zagyva  Formation;
ca. 400 m to the surface; Fig. 2). The samples collected from
the Endrőd and Szolnok formations contained ostracods and
cardiid  molluscs  endemic  to  Lake  Pannon  (for  a  detailed
description  of  the  Pannonian  formations,  see  Juhász  1991;
Juhász et al. 2007; Sztanó et al. 2013a).

Although the Kávás rhinoceros specimens were origi-

nally described as “Lower Pannonian” (this term was tradi-
tionally used for the

fine-grained deep-water deposits

of  Lake  Pannon),  they  were  obviously  recovered  from  the
fluvial  succession  of  the  Zagyva  Formation,  widely  out-
cropping  in  the  vicinity.  The  clayey  sand  embedding  the
bones was deposited in the floodplain of a river that flowed
into  Lake  Pannon  several  tens  of  kilometers  further  to  the
south.

Age assessments within the Pannonian Stage in NW Hun-

gary are based on correlations of biostratigraphic, magneto-
stratigraphic, and seismic stratigraphic data (Magyar et al.
2007), and carry a significant uncertainty. The shelf-margin
slope below Kávás has an estimated age of 8.9 Ma (Magyar
et al. 2007, 2013). The base of the Prosodacnomya zone,
dated to 8.0 Ma in Tihany (Sztanó et al. 2013b), is inferred

Fig. 1. Location map of Kávás, Pannonian Basin, Western Hungary and other Late Miocene Hungarian localities mentioned in the text.

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to be at about 420 m depth in the Nf-3 borehole. Volcanoes
of  the  Little  Hungarian  Plain  Volcanic  Field,  sitting  on  the
eroded  surface  of  the  Zagyva  Formation,  yield  latest
Miocene  and  earliest  Pliocene  age  (e.g.  Ság-hegy,  5.5 Ma;
Wijbrans et al. 2007). The age of the Kávás locality thus can
be  estimated  as  7.0 Ma  (±0.5 Ma),  corresponding  to  the
latest  Tortonian/earliest  Messinian  (i.e.,  Turolian  Land
Mammal Age).

The famous Bérbaltavár (better known as Baltavár: Suess

1861; Pethő 1885; Kormos 1914; Benda 1927; Kretzoi 1985,
1987) mammal locality (MN 12 zone: Bernor et al. 2003,
2005; Kaiser & Bernor 2006) is located some 25 km north-

eastward of Kávás. Seismic profiles between the two loca-
lities show that the Pannonian layers (horizons) are gently
dipping southwards, indicating that Kávás is slightly younger
than Bérbaltavár.

Material and methods

The revised Quaternary time scale of Gibbard et al. (2010)

is used for chronological references in this text. The bottom
and top boundaries of the Pliocene are placed at 5.4 Ma and
2.6 Ma.

The specimens from Kávás collected in 1979 were inven-

toried  as  Rhinoceros  sp.  All  the  specimens  have  the  same
registration  number  V.79.117  in  the  Inventory  Book  of  the
Department of Paleontology and Geology of HNHM. Some
cranial elements (Pandolfi et al. 2015b) have the same regis-
tration number but it is unclear whether cranial and postcra-
nial remains belong to the same individual. Taphonomic data
or photographs of the excavations are not available. However,
similarity  in  dimensions  and  the  existence  of  left  and  right
bones  with  the  same  size  and  shape  suggests  that  the  ele-
ments can belong to a single individual. The postcranial ele-
ments  were  morphologically  compared  with  the  rhinoceros
material collected at several Late Miocene and Pliocene lo-
calities of Eurasia and housed in several museums and insti-
tutions,  as  well  as  with  published  data  (Appendix).  The
anatomical  descriptions  follow  Guérin  (1980)  and  Antoine
(2002), whereas the morphometric approach follows Guérin
(1980).

Institutional Abbreviations:

HNHM, Magyar Természettudományi Múzeum (Hungarian
Natural History Museum), Budapest, Hungary;
IGF, Museo di Storia Naturale, sezione di Geologia e Paleon-
tologia, Florence, Italy;
MFGI,  Magyar  Földtani  és  Geofizikai  Intézet  (Geological
and Geophysical Institute of Hungary), Budapest, Hungary;
MfN, Museum für Naturkunde, Berlin, Germany;
MGGC,  Museo  di  Geologia  Giovanni  Capellini,  Bologna,
Italy;
MNCN,  Museo  Nacional  de  Ciencias  Naturales,  Madrid,
Spain;
MNHN,  Muséum National d’Histoire  Naturelle,  Paris,
France;
MPLBP,  Museo  di  Paleontologia  Luigi  Boldrini  di  Pietra-
fitta, Perugia, Italy;
MPP, Museo di Paleontologia, Università di Parma, Parma,
Italy;
MPUR,  Museo  di  Paleontologia,  Sapienza,  Università  di
Roma, Rome, Italy;
MSNAF,  Museo  di  Storia  Naturale,  Accademia  dei  Fisio-
critici, Siena, Italy;
MSNF, Museo di Storia Naturale, sezione di Zoologia, Flo-
rence, Italy;
NHML, Natural History Museum, London, England;
NHMW, Naturhistorisches Museum, Wien, Austria;
NMB, Naturhistorisches Museum, Basel, Switzerland.

Fig. 2. Stratigraphic position and estimated age of the Kávás out-
crop (asterisk) within the Pannonian sequence. The Neogene suc-
cession at Kávás is based on a hydrocarbon exploration well.

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Systematic palaeontology

Order: Perissodactyla Owen 1848

Family: Rhinocerotidae Gray 1821

Tribe: Rhinocerotini Gray 1821

Genus: Dihoplus Brandt 1878

Type Species: Rhinoceros schleiermacheri Kaup 1832

from the Late Miocene of Eppelsheim, Germany.

“Dihoplus” megarhinus (de Christol 1834)

(Figs. 3—4, Table 1)

Note: Rhinoceros megarhinus de Christol 1834 has some-

times been reported as a synonym of Rhinoceros leptorhinus

Cuvier 1822. However, the Cuvier’s name was based on the
inaccurate description of heterogeneous material belonging
to two or three different taxa (Guérin et al. 1969).

The species Rhinoceros megarhinus was typically as-

signed to the genus Dicerorhinus Gloger 1841 (Guérin 1980,
1982; Guérin & Sen 1998; Guérin & Tsoukala 2013), repre-
sented by the recent species Dicerorhinus sumatrensis (Fisher
1814)  (see  Grooves  1983).  However,  as  noted  by  Pandolfi
(2013)  and  Pandolfi  et  al.  (2015b),  D.  sumatrensis  differs
from  R.  megarhinus  in  having  the  posterior  border  of  the
nasal notch at the level of P2, the dorsal profile of the skull
less concave, the occipital face oblique inclined forward, the
external auditory pseudomeatus open, the protocone and the
hypocone  separated  on  the  upper  premolars  and  the  meta-
cone  fold  well  developed  on  the  upper  premolars  (cranial
material  of  D.  sumatrensis  housed  at  MNHN,  MSNF,

Fig. 3.  “Dihoplus”  megarhinus  from  Kávás.  A  –  atlas  HNHM  V.79.117-7,  anterior  view;  B  –  humerus  HNHM  V.79.117-8,  anterior
view; C – humerus HNHM V.79.117-8, distal view; D – ulna fragment HNHM V.79.117-9, proximal articular surface view; E – radius
HNHM  V.79.117-10,  anterior  view; F  –  radius  HNHM  V.79.117-11,  anterior  view; G  –  radius  HNHM  V.79.117-11,  proximal  view;
H – radius distal epiphysis HNHM V.79.117-12, distal view; I – scaphoid HNHM V.79.117-13, medial view;  L – scaphoid HNHM
V.79.117-13, proximal view; M – magnum HNHM V.79.117-15, anterior view. Scale bars=2 cm.

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NHML  and  NMB:  Pandolfi  2013;  Pandolfi  et  al.  2015b).
The species Rhinoceros megarhinus was recently assigned to
the genus Dihoplus (e.g., Lacombat & Mörs 2008) following
the hypothesis proposed by Heissig (1989, 1996, 1999) who
suggested  an  evolutionary  lineage  leading  from  Dihoplus
schleiermacheri to “Dicerorhinus” megarhinus. Deng et al.
(2011)  ascribed  the  species  R. megarhinus  to  Dihoplus,  but
in the parsimonious trees figured by these authors (Deng et
al.  2011:  fig. S7),  the  genus  Dihoplus  was  paraphyletic  and
D. megarhinus clearly did not form a clade with the species
Dihoplus pikermiensis and Dihoplus ringstroemi. Moreover,
the type species of the genus Dihoplus, D. schleiermacheri,
was  not  included  in  the  analysis  of  Deng  et  al.  (2011).  The
latter species was considered in the unpublished analysis re-
ported by Pandolfi et al. (2014) and Pandolfi (2015a) but it
did not form a clade with D. megarhinus which was included
within the paraphyletic genus Stephanorhinus Kretzoi 1942.
An  assignment  to  the  latter  genus  was  proposed  by  Groves
(1983),  Fortelius  et  al.  (1993)  and  Cerdeño  (1995).  Never-
theless,  de  Christol’s  species  does  not  show  the  typical
morphological characters described as diagnostic for Stepha-
norhinus  (e.g.,  ossified  nasal  septum  or  loss  of  anterior
teeth),  and  the  phylogenetic  relationships  within  this  genus
are  yet  to  be  resolved.  In  agreement  with  Pandolfi  et  al.
(2015b),  we  provisionally  retain  the  species  R.  megarhinus
within the genus Dihoplus.

Material: HNHM V.79.117; 16 post-cranial remains, one

atlas,  one  distal  epiphysis  of  humerus,  one  proximal  frag-
ment  of  ulna,  two  proximal  epiphysis  of  radius,  one  distal
epiphysis  of  radius,  one  damaged  scaphoid,  one  damaged
pyramidal, one fragment of magnum, two fragmentary second
metacarpi,  one  damaged  third  metacarpal,  one  fourth  meta-
carpal, a fragmentary pelvis, one third cuneiform, one proxi-
mal half of fourth metatarsal.

Description and comparison

Atlas: The atlas HNHM V.79.117-7 is poorly preserved

and the transverse processes are partially damaged (Fig. 3A).
In dorsal view,

the dorsal tubercle is relatively large, the two

alar  foramina  are  partially  damaged  but  appear  large.  The
posterior border of the bone is concave. In ventral view, the
ventral  tubercle  is  developed  and  extends  posteriorly.  In
anterior  view,  two  deep  articular  surfaces  for  the  occipital
condyles are present (Fig. 3A); they are externally delimited
by  a  marked  edge.  In  posterior  view,  the  articular  surfaces
for the axis are flat.

In the atlas of D. schleiermacheri from Eppelsheim

(NHML  1284)  the  anterior  articular  surfaces  are  more  dis-
tally  separated  than  in  the  studied  specimen.  In  anterior
view, the atlas of S. etruscus (specimens from Capitone and
Upper  Valdarno:  Appendix)  differs  from  the  studied  speci-
men in having a more rounded proximal-lateral border of the
articular surface and in being less massive. A relatively well
preserved  atlas  of  “D.”  megarhinus  from  Montpellier
(NHMB Mp922) displays the same morphological characters
described for the atlas from Kávás. Unfortunately, no atlas of
“S.” miguelcrusafonti is known, and we had no opportunity

to observe directly any atlas attributed to S. jeanvireti or
D. pikermiensis.

Humerus: Only a distal epiphysis of a humerus

V.79.117-8  is  kept  at  HNHM.  The  trochlea  is  anteriorly
damaged (Fig. 3B—C). In anterior view, the medial border of
the medial lip of the trochlea is slightly convex whereas the
lateral  border  of  the  lateral  lip  is  straight  (Fig. 3B—C).
The lateral tuberosity is well developed and large, the medial
tuberosity  is  much  smaller.  The  lateral  epicondylar  crest  is
relatively  short,  marked  and  well  developed.  The  trochlear
fossa is transversally elongated. In posterior view, the lateral
epicondylar  crest  is  well  developed,  the  olecranon  fossa  is
wide  and  deep.  The  lateral  epicondyle  is  large  and  well
developed  and  the  medial  epicondyle  is  massive.  In  distal
view, the medial lip of the trochlea has a convex medial bor-
der; the posterior border of the trochlea is regularly concave
and  the  medial  epicondyle  extends  posteriorly  (Fig. 3B—C).
The lateral tuberosity is well developed and rounded.

In distal view, the humeri of D. pikermiensis (Appendix)

display  a  smaller  and  less  anterior-posteriorly  developed
lateral tuberosity. The studied specimen has a larger olecra-
non  fossa  than  in  S.  jeanvireti  (Appendix)  and  the  bone  is
more massive than in S. jeanvireti and S. etruscus (Appendix).
No  morphological  data  are  available  on  the  humeri  of
“S.” miguelcrusafonti and D. schleiermacheri. The humeri of
“D.” megarhinus from Monte Giogo (MPP: Simonelli 1897)
and Val di Pugna (MSNAF 7100) have a marked antero-pos-
terior crest in the lateral side of the distal epiphysis, a large
olecranon  fossa  and  massive  epicondyles  as  in  the  studied
specimen.  The  humerus  of  “D.”  megarhinus  from  Saint-
Laurent (Guérin et al. 1969: figs. 18—19) displays a sinuous
medial border of the medial lip of the distal trochlea and the
posterior-medial  epicondyle  is  massive  as  well  as  in  the
specimen  from  Kávás.  The  specimens  of  “D.”  megarhinus
from  Montpellier  (Appendix)  share  several  morphological
characters  with  those  from  Kávás:  e.g.,  the  posterior-lateral
epicondyle on the humerus is well developed and larger than
the medial one, the lateral epicondylar crest is well evident.
The dimensions of the distal epiphysis of the humerus from
Kávás  are  slightly  larger  than  those  of  “D.”  megarhinus
from several Pliocene localities (Table 1).

Ulna: The ulna is represented by a fragment of a proximal

epiphysis HNHM V.79.117-9 (Fig. 3D). In anterior view, the
medial and lateral sides of the articular surface for the hu-
merus are concave. The articular surfaces for the radius are
not evident due to the

bad preservation. The sigmoid incisure

is relatively high, flat, and distally delimited by a deep fora-
men (Fig. 3D).

The ulna of D. schleiermacheri differs from the studied

specimen  in  having,  in  anterior  view,  a  more  asymmetrical
articular surface (Guérin 1980: fig. 34C). With respect to the
specimen  from  Kávás,  the  proximal  articular  surface  of  the
ulna of D. pikermiensis (Appendix) is more transversally de-
veloped  and  the  medial  lip  is  more  concave.  The  proximal
articular  surface  of  the  ulna  of  S.  jeanvireti  (Appendix)  ap-
pears  to  be  more  slender  than  that  of  the  studied  specimen
and the medial lip is higher and more concave. In S. etruscus
(Appendix) the sigmoid incisure is shorter than in Kávás and
the  proximal  articular  surface  is  more  symmetrical  in  ante-

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rior view. The morphological characters of the studied speci-
men are also evident in a fragmentary ulna of “D.” megarhi-
nus from Montpellier (NMB Mp1008).

Radius: Two proximal halves of radius, HNHM

V.79.117-10 and HNHM V.79.117-11 (Fig. 3E—G), and
a fragmentary distal epiphysis, HNHM V.79.117-11, are pre-
served (Fig. 3H).

In the proximal epiphysis, in anterior view, the coronoid

process  is  prominent,  the  bicipital  tuberosity  is  slightly  de-
pressed,  and  the  posterior  proximal  apophysis  is  evident
(Fig. 3E—F). In posterior view, a long and very narrow medial
articular  surface  for  the  radius  is  present  on  the  specimen
HNHM  V.79.117-10  whereas  it  is  absent  on  HNHM
V.79.117-11.  A  larger,  slightly  concave  and  subtriangular
lateral articular surface for the radius is present on both spe-
cimens. This surface is slightly smaller and laterally delimi-
ted by a marked groove on HNHM V.79.117-11.

In proximal view, the medial and lateral articular surfaces

are  subquadrangular  (Fig. 3G).  The  medial  border  of  the
proximal  surface  is  convex,  whereas  the  anterior  border  is
concave  only  at  the  level  of  the  coronoid  apophysis
(Fig. 3G).  The  lateral  border  is  straight  and  oblique  on
HNHM  V.79.117-11.  The  angle  between  the  posterior  bor-
der of the medial articular surface and that of the lateral one
is obtuse.

On the distal epiphysis of HNHM V.79.117-12, in anterior

view, the articular surface for the scaphoid is partially
observable, whereas the medial and lateral styloid processes
are not preserved (Fig. 3H). The distal border of the articular
surface for the semilunar is convex. In distal view, the distal
articular surface is well developed, its posterior medial por-
tion extends backwards. The anterior border of the epiphysis
is concave at the level of the extensor carpi radialis. The ar-
ticular surface for the semilunar is concave whereas that for
the scaphoid is anteriorly concave and posteriorly convex.

Compared with the studied material, “S.” miguelcrusa-

fonti  from  the  Pliocene  of  Spain  displays  a  rounded  proxi-
mal-medial  articular  surface  on  the  radius  and  a  less  deve-
loped posterior apophysis on the proximal epiphysis (Guérin
&  Santafé-Llopis  1978:  pl. 5A—B).  The  studied  specimens
differ  from  the  Pliocene  S.  jeanvireti  (Appendix),  which
displays,  in  proximal  view,  a  less  obtuse  angle  between
the  posterior  borders  of  the  medial  and  lateral  articular
surfaces. The remains of the latest Pliocene—Early Pleistocene
S.  etruscus  (Appendix)  are  smaller  than  those  collected
at  Kávás,  the  proximal  lateral  surface  on  the  proximal
epiphysis of the radius is less developed and its lateral  bor-
der is convex. In  D. pikermiensis (Appendix) the proximal-
lateral  articular  surface  for  the  ulna  is  slightly  more
developed,  the  anterior  border  of  the  proximal  epiphysis
has a concavity in the middle, the proximal-lateral articular
surface  for  the  humerus  extends  forward  and  its  anterior
border is at the same level than that of the proximal-medial
surface.  The  radius  of  “D.”  megarhinus  from  Montpellier
(Appendix) shares with the specimens from Kávás a convex
medial  border  of  the  proximal  epiphysis  and  a  concave
anterior  border  at  the  level  of  the  coronoid  apophysis.
In  some  specimens  from  Montpellier  the  posterior-medial
articular  surface  for  the  ulna  is  not  evident.  The  distal
epiphysis  of  a  radius  from  Val  di  Pugna  (MSNAF  4754)
is very damaged, but resembles the specimens from Kávás in
having, in distal view, a straight lateral border of the articular
surface and  a slightly concave posterior border of the lateral
half;  moreover,  the  external    tuberosity  of  the  anterior  face
is  rounded  and  large.  These  features  are  also  more  evident
in  “D.”  megarhinus  from  Montpellier  than  in  any  other
Pliocene species (Guérin 1972: fig. 2B). The dimensions of
the  proximal  epiphyses  from  Kávás  are  slightly  larger  than
those  of  “D.”  megarhinus  from  several  Pliocene  localities
(Table 1).

Table

1: Measurements (in mm) of the postcranial remains of “Dihoplus” megarhinus from Kávás (Zala subbasin, Hungary), compared

with those of “D.” megarhinus from Montpellier (Early Pliocene, France), Udunga (latest Pliocene, Russia), and with the minimal and
maximal values given by Guérin (1980). DTD – distal transverse diameter; DAPD – distal antero-posterior diameter; DTDth – distal
transverse diameter of the trochlea ; TDof – transverse diameter of the olecranon fossa; PTD – proximal transverse diameter;
PAPD – proximal antero-posterior diameter; TD – transverse diameter; l

×××××H – breadth and height; L – length; DTDmax – maximal

distal transverse diameter.

Measurement Kávás

(HNHM

v.79.117)

Montpellier
(NMB various
specimens)

Western Europe
(from Guérin, 1980)

Udunga
(from Fukuchi et al., 2009)

Humerus DTD

171

153.44–160.16

144–166

154.4–158.9

Humerus DAPD

137.2

125.1–132.96

100–135

99.5–120.8

Humerus DTDth

120.1

98.61–126.43

Humerus TDof

53.5

43.65–55.76

Radius PTD

120.4

108.13–111.19

94.5–116

96.1–110.6

Radius PAPD

85–86.1

70.52–80.10

61–82

66.8–82

Scaphoid TD

58.7

61.6

50–72.5

47.5–65.1

Magnum TD

44.5–61

51.1–60.4

Pyramidal l

×H

54.7

×63

53.64

×65.75

49–65.5

×53–64

41.9–65.6x52.9–71.6

MCII PTD

55.9–59.3

53.46–57.52

39–58.5

49.5–58.1

MCIII PAPD

50.5

53.98–57.22

48–58

44.5–58.2

MCIV PTD

48.5

53.02

48–54

42–67.3

MCIV PAPD

46.1

44.63

36.5–47

39.5–53.2

Third cuneiform l

×H×L 57.1×29.9×57.7

48.5–57

×29.5–35×47–61 51.3–60.6×28–35.2×54.7–66.6

MTIV L

167.5

164–182

MTIV DTDmax

40.5

39–42

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Scaphoid: The scaphoid HNHM V.79.117-13 is partially

damaged on its anterior-proximal border (Fig. 3I—L). In me-
dial view, the posterior border of the bone is straight whereas
the  anterior  one  is  convex  in  its  proximal  half  (Fig. 3I).  In
lateral view, the bone is very damaged, the articular surface
for the semilunar is not evident. In the same view, the distal
articular surfaces are evident. They are composed by a small
anterior  articular  surface  for  the  semilunar  connected  with
a  larger  one  for  the  magnum.  In  dorsal  view,  the  articular
surface  is  anteriorly  damaged  and  transversally  covers  the
proximal face of the bone.

The scaphoid of D. schleiermacheri from Eppelsheim

(NHML  1281)  differs  from  the  studied  specimen  in  being
shorter  and  in  having,  in  medial  view,  a  convex  posterior-
distal border. The scaphoid of “S.” miguelcrusafonti differs
from  that  of  Kávás  in  having,  in  medial  view,  a  larger  and
higher  distal  articular  surface  (Guérin  1980:  fig. 63B).  In
S.  jeanvireti  the  anterior  tuberosity  is  more  marked  and
developed  and  the  proximal  articular  surface  is  not  evident
in  medial  view.  The  scaphoid  of  S.  etruscus  is  shorter  and
appears massive. The scaphoid from Kávás resembles those
of “D.” megarhinus from Montpellier in the morphology of
the  anterior  tuberosity,  of  the  posterior  border  of  the  bone
and  in  the  development  of  the  distal  articular  surface.  The
unique  dimension  obtained  from  the  scaphoid  falls  within
the dimensional range of “D.” megarhinus (Table 1).

Pyramidal: The pyramidal HNHM V.79.117-14 is very

damaged  and  only  the  lateral  half  of  the  bone  is  well  pre-
served. In anterior view, the proximal-lateral border is con-
vex,  whereas  the  lateral-distal  one  is  concave.  A  relatively
large tuberosity is present on the lateral border. In proximal
view,  the  proximal  articular  surface  is  concave  anterior-
posteriorly  and  convex  lateral-medially.  In  distal  view,  the
distal articular surface is flat and subtrapezoidal.

The distal articular surface on the pyramidal of “S.” miguel-

crusafonti  (Appendix)  is  smaller  with  rounded  angles;  this
surface  is  more  rounded  in  S.  jeanvireti  than  in  the  studied
specimen. The pyramidal of S. etruscus is smaller and shorter
than  the  studied  specimen.  Unfortunately,  pyramidals  cer-
tainly  attributable  to  D.  schleiermacheri  or  D.  pikermiensis
have not been found in the visited collections and any useful
morphological character cannot be obtained from the figures
published  by  Guérin  (1980:  fig. 36E).  The  pyramidal  of
“D.”  megarhinus  from  Montpellier  displays  the  same  mor-
phology described for the Kávás specimen. The dimensions
of  the  pyramidal  fall  within  the  dimensional  range  of
“D.” megarhinus (Table 1).

Magnum: Only the anterior face of the magnum HNHM

V.79.117-15 is preserved (Fig. 3M). In anterior view, the an-
terior face of the bone is pentagonal. The distal border is
convex, and the medial border has a slight concavity in its
distal half (Fig. 3M).

The magnum of “S.” miguelcrusafonti (Appendix) is very

damaged; the preserved portion of the anterior face appears
less massive than in the studied specimen. The anterior face
of the magnum of S. jeanvireti is proportionally higher and
transversally shorter than that from Kávás whereas the mag-
num of S. etruscus is generally smaller. The magnum of
“D.” megarhinus from Montpellier is massive and displays

a well developed anterior tuberosity as in the specimen from
Kávás. The unique dimension obtained from the magnum
falls within the dimensional range of “D.” megarhinus
(Table 1).

Second Metacarpal: In proximal view, the articular sur-

face  for  the  trapezoid  of  the  second  metacarpal  HNHM
V.79.117-16 has rounded angles (Fig. 4A); it is larger in its
anterior half than in the posterior one. A developed tuberosity
occurs  at  the  posterior  end  of  the  proximal  epiphysis
(Fig. 4A). In lateral view, the articular surface for the mag-
num is long; its proximal border is not regularly convex and
it is separated from the proximal articular surface by a marked
edge.  The  articular  surface  for  the  third  metacarpal  (small,
narrow and long) is separated from that for the magnum by
a  very  bland  edge.  The  distal  border  of  the  lateral  articular
surface  is  concave  and  it  is  delimited  by  a  marked  groove.
On the specimen HNHM V.79.117-17 the proximal articular
surface  is  rounded,  concave  lateral-medially  and  slightly
convex  anterior-posteriorly;  it  is  transversally  elongated  in
its anterior half but does not reach the lateral and medial bor-
ders of the proximal epiphysis (

Fig. 4A—B

In D. schleiermacheri the lateral articular surface on the

proximal  epiphysis  is  separated  in  two  facets  by  a  bland
groove and, in proximal view, the proximal articular surface
is  more  rounded  (Guérin  1980:  fig. 39B).  The  MCIIs  of
D.  pikermiensis  differ  from  the  studied  specimen  in  having
a  less  developed  anterior-lateral  tuberosity,  a  much  wider
proximal articular surface and a flat proximal-lateral articu-
lar  surface.  The  proximal  articular  surface  on  the  MCII  of
“S.”  miguelcrusafonti  from  Layna  (MNCN)  is  narrower
whereas  the  lateral  articular  surface  is  more  concave  proxi-
mal-distally.  The  specimen  from  Kávás  differs  from  the
Pliocene  S.  jeanvireti  which  displays  a  flat  proximal-lateral
articular  surface  and  from  S.  etruscus  which  displays  little
developed  medial  and  lateral  tuberosities  on  the  proximal
epiphysis. The MCIIs of “D.” megarhinus from Montpellier
share  several  morphological  characters  with  those  from
Kávás: e.g., the proximal-lateral articular surface is concave
and  the  anterior-medial  tuberosity  is  well  evident  in  proxi-
mal  view.  The  dimensions  of  the  MCIIs  from  Kávás  are
close to the maximal values of “D.” megarhinus (Table 1).

Third Metacarpal: The proximal half of the third meta-

carpal  HNHM  V.79.117-18  is  badly  preserved  (Fig. 4C—E).
In anterior view, the proximal-medial tuberosity is prominent
and  the  proximal  border  of  the  proximal  articular  surface  is
concave. In proximal view the articular surface for the mag-
num  is  subtrapezoidal,  with  a  convex  medial  border,  and
a  slightly  concave  anterior  border  (Fig. 4C).  This  surface  is
separated from that for the uncinate, smaller and subtriangu-
lar, by a strong saliency. In lateral view, the anterior-proximal
and  the  posterior  articular  surfaces  are  well  separated  by
a  marked  groove  (Fig. 4E).  The  anterior-proximal  surface  is
subtrapezoidal and is proximally joined with that for the unci-
nate. The posterior surface is subelliptical, with the maximal
axes parallel to the posterior border of the diaphysis.

The studied material differs from D. schleiermacheri in

which the posterior-lateral articular surface on the MCIII is
rounded and larger than the anterior-lateral one and the pro-
ximal articular surface on the MCIII is less developed trans-

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versally (Appendix). In D. pikermiensis (Appendix) the pos-
terior-lateral  articular  surface  is  wider  and  well  developed,
the  articular  surface  for  the  uncinate  is  less  developed  and
the medial border of the proximal epiphysis is less expanded
anterior-posteriorly.  Compared  with  the  studied  material
“S.”  miguelcrusafonti  displays  a  less  developed  proximal-
medial tuberosity on MCIII (Guérin & Santafé-Llopis 1978).
The  studied  specimen  differs  from  S.  jeanvireti  which  has
a convex anterior border and a subtriangular lateral articular
surface. The MCIII of S. etruscus differs from that of Kávás
in having a small and subelliptical proximal-lateral articular

surface  and  a  well  developed  and  subtriangular  posterior-
lateral surface. The MCIIIs of “D.” megarhinus from Mont-
pellier  share  several  characters  with  the  studied  specimen:
the  anterior  border  of  the  proximal  epiphysis  is  usually
slightly concave but is also straight in some specimens, the
posterior-lateral articular surface is  subelliptical but appears
slightly wider than that from Kávás.

Fourth Metacarpal: On the fourth metacarpal HNHM

V.79.117-19, in proximal view, a broad articular surface for
the uncinate is present (Fig. 4F—G). The latter surface is sub-
triangular, its medial border is slightly convex, whereas the
anterior one is straight. In medial view, the two articular sur-
faces  for  the  third  metacarpal  are  badly  preserved  and  only
a rather rounded posterior one is evident. In posterior-lateral
view, the proximal articular surface slightly extends over the
posterior-lateral border of the proximal epiphysis giving two
small,  elongated  and  narrow  surfaces,  distally  delimited  by
two  marked  depressions.  MCIV  of  D.  schleiermacheri  has
never been reported from Eppelsheim (Guérin 1980).

In respect to the studied specimen, the proximal epiphysis

of D. pikermiensis and S. jeanvireti are transversally longer
and  anterior-posteriorly  shorter;  both  species  display,  how-
ever,  a  different  shape  of  the  proximal  epiphysis.  In  “S.”
miguelcrusafonti  the  posterior  border  of  the  proximal  epi-
physis  displays  a  marked  groove  and  the  proximal  articular
surface  is  less  developed  than  the  proximal  epiphysis
(Guérin 1980: fig. 71B). The proximal epiphysis of S. etrus-
cus, in proximal view, is rather similar to that of the studied
specimen but its posterior border is generally straight and the
lateral articular surface is partially evident. The shape of the
proximal epiphysis of the specimen from Kávás is similar to
that  of  the  MCIVs  of  “D.”  megarhinus  from  Montpellier.
The  dimensions  of  the  MCIV  from  Kávás  fall  within  the
values of “D.” megarhinus (Table 1).

Pelvis: The pelvis is represented only by a fragment of

acetabulum and ischium HNHM V.79.117-20. The proximal
border of the acetabulum is regularly convex, whereas the
posterior-proximal border is straight (Fig. 4H). The pre-
served portion of the articular cavity is deep and surrounded
by a sharp edge.

The posterior-proximal border of the acetabulum from

Montpellier is straight as well as that from Kávás and similar
to the specimen from Rio Secco (MGGC 9350). S. etruscus
differs from the studied specimen in being smaller and in
having a more rounded acetabulum. In S. jeanvireti the angle
between the dorsal border of the acetabulum and the dorsal
border of the ischium is more obtuse.

Third Cuneiform: In anterior view, the anterior face of

the  third  cuneiform  HNHM  V.79.117-21  is  rectangular
(Fig. 4I—L).  The  proximal  border  is  slightly  concave  on  its
medial half and slightly convex in its lateral half. The distal
border is slightly convex. The medial and lateral borders are
straight, and the angle between the distal border and the me-
dial one is approximately of 90°. In proximal view, the pro-
ximal articular surface is wide and subtriangular. The medial
face of the bone is badly preserved; the anterior and posterior
articular surfaces for the second metatarsal are subsquare and
the anterior one is slightly higher. In posterior-lateral view,
two  articular  surfaces  are  present.  The  posterior-proximal

Fig. 4.  “Dihoplus”  megarhinus  from  Kávás.  A  –  MCII  HNHM
V.79.117-16,  proximal  view;  B  –  MCII  HNHM  V.79.117-16,
anterior  view;  C  –  MCIII  HNHM  V.79.117-18,  proximal  view;
D  –  MCIII  HNHM  V.79.117-18,  anterior  view;  E  –  MCIII
HNHM V.79.117-18, lateral view; F – MCIV HNHM V.79.117-19,
proximal  view;  G  –  MCIV  HNHM  V.79.117-19,  anterior  view;
H – pelvis HNHM V.79.117-20, acetabular view; I – third cunei-
form  HNHM  V.79.117-21,  proximal  view;  L  –  third  cuneiform
HNHM V.79.117-21, anterior view; M – MTIV HNHM V.79.117-22,
proximal  view;  N  –  MTIV  HNHM  V.79.117-22,  anterior  view.
Scale bars=2 cm.

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one is elliptical whereas the anterior-distal one is triangular,
wide and flat.

The anterior face of the third cuneiform in D. schleierma-

cheri is proportionally higher and transversally shorter than
that  of  the  studied  specimen  (Guérin  1980:  fig. 50C).  The
third cuneiform of “S.” miguelcrusafonti is rather similar to
the studied specimen but, in distal view, the anterior-medial
side  of  the  face  appears  less  developed  anterior-posteriorly.
The  studied  specimen  differs  from  S.  jeanvireti  which  dis-
plays a concave lateral border and a convex medial border of
the  anterior  face.  The  dorsal  border  of  the  anterior  face  in
S.  etruscus  is  more  concave,  whereas  the  distal  border  is
more  convex  than  in  the  studied  specimen;  moreover  the
anterior face in S. etruscus appears higher and transversally
shorter.  The  dimensions  of  the  third  cuneiform  from  Kávás
fall within the values of “D.” megarhinus (Table1).

Fourth Metatarsal: In anterior view, the medial border of

the  fourth  metatarsal  HNHM  V.79.117-22  is  sinuous
(Fig. 4M—N).  The  insertion  of  the  muscle  interossei  is  long
and  reaches  the  distal  half  of  the  diaphysis.  In  proximal
view, the articular surface is rounded, its posterior border has
a  concavity  in  the  middle  and  is  delimited  by  a  marked
groove  (Fig. 4M).  The  two  articular  surfaces  for  the  third
metatarsal, on the medial face of the bone, are of about the
same  size;  the  anterior  one  is  subtrapezoidal,  whereas  the
posterior  one  is  rounded.  These  two  surfaces  are  separated
by a marked groove. Moreover, the anterior surface is proxi-
mally joined with the proximal articular surface.

The MTIV of D. schleiermacheri displays, in proximal

view,  a  well  developed  posterior  tuberosity  and,  in  medial
view, the posterior articular surface for the third metatarsal is
joined  with  the  proximal  articular  surface  (Guérin  1980:
fig. 55E). In D. pikermiensis the proximal articular surface is
rounded  and  the  two  medial  articular  surfaces  are  partially
evident in proximal view. In respect to the studied specimen,
the proximal articular surface of S. jeanvireti is less rounded
and  less  developed  than  the  proximal  epiphysis.  The  proxi-
mal articular surface in S. etruscus is triangular and smaller
than that of the specimen from Kávás. In  “D.” megarhinus
from  Montpellier,  as  in  the  studied  specimen,  the  proximal
articular  surface  is  rounded,  the  anterior  medial  surface  for
the third metatarsal is joined with the proximal epiphysis and
the  posterior  medial  surface  is  rounded  (Guérin  1972:
fig. 20B).  The  dimensions  of  the  MTIV  from  Kávás  fall
within the values of “D.” megarhinus (Table 1).

Discussion and conclusions

Although there are numerous localities with Turolian land

mammal  remains  from  Hungary,  remains  of  land  mammals
tend  to  be  sparse.  However,  a  few  localities  contain  speci-
men-rich  land  mammal  assemblages  dominated  by  large-
sized  mammals.  Kretzoi  (1982)  gave  a  detailed  list  of  the
so-called  Hipparion-fauna localities from the Late Miocene
of  the  Carpathian  Basin  and  sketched  the  biochronological
correlation  among  the  most  important  localities  (Kretzoi
1982,  1985,  1987).  Kretzoi  established  the  Sümegium  and
the  Hatvanium  (these  two  stages  approximately  correspond

to  the  MN 12)  and  the  Bérbaltavárium  (approximately  cor-
responds  to  the  MN 13;  Bérbaltavár  is  the  recent  name  of
a  small  village  that  was  called  Baltavár  earlier).  He  placed
Baltavár  and  Polgárdi,  the  two  faunistically-richest  late
Turolian  land  mammal  localities  of  Western  Hungary,  into
the  Bérbaltavárium.  This  correlation  was  used  later  by
Kordos  (1992)  and  Gasparik  (2001).  Kretzoi  (1983)  also
sketched a biostratigraphic chart using Hipparion species as
biostratigraphic-index  forms.  However,  Kaiser  &  Bernor
(2006)  revised  the  Baltavár  “hipparions”  and  pointed  out
that  Baltavár  is  older  than  was  believed  earlier.  Its  age  is
MN 12 rather than MN 13 and Polgárdi belongs to MN 12 or
MN 13. This result fits well with older opinions because Bal-
tavár  assemblages  are  compositionally  very  similar  to  the
world-famous  Pikermi  fauna.  A  similar  dating  has  been
inferred  by  Gasparik  (2004)  on  the  basis  of  proboscidean
material:  MN 12  for  Baltavár  and  MN 13  for  Polgárdi.  The
proboscidean  record  from  Baltavár  is  still  under  revision
because  the  two  species  that  have  been  described  here  (cf.
Tetralophodon  longirostris  Kaup  1832  and  cf.  Mammut
borsoni  Hays  1834)  show  some  characteristics  which  indi-
cate  that  these  specimens  must  probably  be  reassigned  to
other  species  [Tetralophodon  atticus  (Wagner  1857)  and
Mammut obliquelophus (Mucha 1980)], as was suggested by
Markov (2008).

Latest Miocene (MN 12 or 13) rhinoceroses are poorly

documented in Western Hungary and are represented by rare
remains.  As  far  as  the  rhinocerotid  remains  from  the  above
mentioned localities are concerned, “Dihoplus” megarhinus
was not described from any of them, but two other species,
identified  as  Aceratherium  incisivum  and  D.  schleierma-
cheri,  were  found  (Kretzoi  1952,  1982;  Kordos  1992).  The
latter  species  has  been  reported  at  Baltavár  (MN 12)
(Rhinoceros  pachygnathus  in  Pethő  1885;  Giaourtsakis
2009).  However,  a  fragment  of  hemimandible  with  p4-m3
(L.sz.Ob-331)  housed  at  the  Geological  Museum  of  the
MFGI displays morphological characters (a short paralophid
and  a  mesial-lingual  cingulum)  that  suggest  a  similarity  to
Aceratherium. Some specimens housed at the MFGI can be
ascribed to Aceratherium sp. (an isolated DP4 v13.00339.1;
an  isolated  lower  molar  v13.00335.1)  or  Rhinocerotidae
indet. (an isolated and much worn M3 v13.00376.1) whereas
only  a  calcaneum  (v13.00340.1)  and  perhaps  a  fragment
of  juvenile  mandible  with  dp1-dp2  can  be  identified  as
Dihoplus. Nevertheless, the calcaneum (v13.00340.1) differs
from  D.  schleiermacheri  in  having  a  more  developed  tuber
calcanei  and,  in  posterior  view,  a  clearly  evident  articular
surface  for  the  cuboid.  Moreover,  the  values  of  the  trans-
verse diameter of the sustentaculum tali (DT=98 mm) and of
the  anterior-posterior  diameter  of  the  tuber  calcanei
(DAP=88 mm) are larger than those reported for D. schleier-
macheri  by  Guérin  (1980)  and  are  close  to  the  maximal
values of “D.” megarhinus (Guérin 1980: tab. 108). Unpub-
lished remains of a rhinoceros housed at the NMB have been
collected  at  Polgárdi  and  include  indeterminable  fragments
of  teeth  and  fragments  of  bones.  Among  the  other  remains,
a  damaged  proximal  epiphysis  of  radius  morphologically
resembles the specimens from Kávás and can be assigned as
“D.” cf. megarhinus.

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“D.” megarhinus has usually been documented in the

Pliocene deposits (Guérin et al. 1969; Guérin 1980; Pandolfi
2013), but its presence has been also suggested in latest Mio-
cene,  MN 13  (late  Messinian),  localities  of  Baccinello  V3
(Toscana,  Italy:  Hürzeler  &  Engesser  1976;  Pandolfi  2013;
Pandolfi et al. 2015b) and Monticino Quarry (Emilia-Romagna,
Italy: De Giuli 1989; Pandolfi 2013; Pandolfi et al. 2015b).
The  record  of  Kávás,  which  is  older  than  the  above  men-
tioned  Italian  records,  strongly  reinforces  the  occurrence  of
“D.”  megarhinus  during  the  Miocene  in  Europe  and  sug-
gests a critical revision of several findings usually identified
as Dihoplus  sp.  (Novo  Elisavetovka,  Ukraine,  MN 12:  Ale-
xejewa  1916;  Giaourtsakis  et  al.  2006;  Pandolfi  et  al.
2015b),  Stephanorhinus  sp.  (Moncucco,  Northern  Italy,
MN 13: Angelone et al. 2011) or Dihoplus schleiermacheri?
(Verduno, Northern Italy, MN 13: Colombero et al. 2014).

Moreover, the occurrences of “D.” megarhinus through-

out the latest Miocene suggest that this species spread from
the  Pannonian  Basin  towards  Italy  during  the  MN 13.  The
latter hypothesis is also supported by the dispersal pattern of
the genus Hippotherium recently suggested by Bernor et al.
(2011).  According  to  these  authors,  Hippotherium  is  not
documented  from  the  Baccinello  area  until  the  base  of  the
MN 13 and it may have emigrated from the Pannonian area.

The occurrences of “D.” megarhinus in Western Hungary

(MN 12  and  13)  and  Italy  (MN 13)  also  suggests  a  biogeo-
graphic segregation of Rhinocerotini species in Europe during
the  latest  Miocene.  In  fact,  D.  schleiermacheri  is  the  sole
Rhinocerotini  species  in  Western  Europe  during  the  latest
Miocene (Guérin 1980; Cerdeño, 1992; Heissig 1996, 1999)
whereas  D.  pikermiensis  and  C.  neumayri  represented  the
two  rhinocerotine  species  during  the  Turolian  (late  Torto-
nian-Messinian,  approximately  9-5.3 Ma)  in  Southeastern
Europe  (Geraads  1988;  Geraads  &  Spassov  2009).
D.  schleiermacheri,  D.  pikermiensis  and  C.  neumayri  be-
came extinct at the end of the Miocene (Guérin 1980; Heissig,
1996, 1999) whereas “D.” megarhinus occurred in southern
France  (MN 14),  in  Turkey  (MN 15)  and  elsewhere  too  in
Europe (Guérin 1980; Radulescu & Samson 1985; Guérin &
Sen 1998; Pandolfi 2013).

Acknowledgements:  We  are  grateful  to  E.  Cerdeño,  V.
Codrea  and  the  editor  A.  Tomašových  for  their  useful
comments  and  suggestions.  LP  thanks  E.  Cioppi  (IGF),
E.  Bodor  (MFGI),  O.  Hampe  (MfN),  C.  Sarti  (MGGC),
P.  Pérez  Dios  (MNCN),  M.C.  De  Angelis  (MPLBP),
R. Manni (MPUR), F. Farsi (MSNAF), P. Brewer (NHML),
U.  Göhlich  (NHMW)  and  L.  Costeur  (NMB)  for  their  help
and  assistance  during  his  visits  to  the  rhinoceros  fossil  col-
lections.  LP  also  thanks  L.  Maiorino  for  pictures  of  the
specimens housed at MNHN and MPP. LP thanks the Euro-
pean

Commission’s

Research

Infrastructure

Action,

EU-SYNTHESYS  project  AT-TAF-2550,  DE-TAF-3049,
GB-TAF-2825,  HU-TAF-3593,  ES-TAF-2997;  part  of  this
research  received  support  from  the  SYNTHESYS  Project
http://www.synthesys.info/ which  is  financed  by  European
Community  Research  Infrastructure  Action  under  the  FP7
“Capacities”  Program.  This  is  MTA-MTM-ELTE  Paleo
contribution No. 212.

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Appendix

Source for postcranial comparison material. Institutional abbreviations are reported in the text. nc = no code.

Bibliographic Source

“Dihoplus” megarhinus
Locality: Montpellier, Saint-Laurens, Monte Zago, Rio Secco,
Val di Pugna-Fangonero.
References: Simonelli (1897); Guérin et al. (1969); Guérin
(1980); Pandolfi (2013).
“Stephanorhinus” miguelcrusafonti
Locality: Layna, Perpignan.
References: Guérin & Santafé-Llopis (1978).
Stephanorhinus jeanvireti
Locality: Vialette, Villafranca d’Asti.
References: Guérin (1972, 1980).
Stephanorhinus etruscus
Locality: Senèze, Capitone, Upper Valdarno
References: Ambrosetti (1972); Guérin (1972, 1980);
Pandolfi & Petronio (2011).
Dihoplus schleiermacheri
Locality: Eppelsheim.
References: Guérin (1980).

Direct Observations

HNHM
Stephanorhinus jeanvireti
Ajnácsko: Humerus b.801; Pyramidal b.918, b.828;
MC2 b.807; MC4 b.806.

IGF
Stephanorhinus jeanvireti
Montopoli: two anterior and two posterior limbs 1075.
Stephanorhinus etruscus
Upper  Valdarno:  Assembled  skeleton  716,  3098,  2293v;
Almost  complete  anterior  limb  731;  Humerus  730,  717,
2209v,  14840,  488v;  Radius  2211v,  2212v,  2214v,  4566v,
4567v, 488v; Ulna 4566v, 4567v; MC2 1355v; MC3 1355v,
2231v, 488v; MC4 2232v, 488v; MT4 2233v, 487v.

MFGI
Stephanorhinus jeanvireti
Pula: Radius v.18511, v.18514, v.18515.
Dihoplus schleiermacheri
Pannonian Basin: Radius v.11117; MC3 v.11110.

MfN
Dihoplus schleiermacheri
Eppelsheim: Radius MbMa28312; MC3 MbMa28313,
MbMa 28307.
Dihoplus pikermiensis
Pikermi: Humerus MbMa24815, MbMa24817; MC2
MbMa28279; MC3 MbMa28281, MbMa28283.

MGGC
“Dihoplus” megarhinus
Rio Secco: Pelvis 9350.

Montegiogo: Atlas 9372.
Monte Zago?: Complete anterior limb (cast) nc.
Stephanorhinus jeanvireti
Monte Pastore: Humerus nc.

MNCN
Dihoplus schleiermacheri
Venta del Moro: Third Cuneiform 11817.
“Stephanorhinus” miguelcrusafonti
Layna:  Pyramidal  23785;  Magnum  23783;  MC2  70374;
MT4 23767.
Stephanorhinus etruscus
Cullar  de  Baza  I:  Scaphoid  13141;  Pyramidal  13143;  MC2
13144.
El-Rincon:  Humerus  41874;  Radius  41869;  MC3  41870;
MT4 41871.
Huéscar: Humerus 19206; MC3 55139; Pelvis 19207.
La Puebla de Valverde: MC2 32751; MC4 54888.

MPLBP
Stephanorhinus etruscus
Pietrafitta: Two almost complete mounted skeletons.

MPUR
Stephanorhinus etruscus
Capitone: Almost complete skeleton 1500.

MSNAF
“Dihoplus” megarhinus
Val di Pugna-Fangonero: Humerus 7100; Radius 4754.
Stephanorhinus etruscus
Castelnuovo di Barardenga Scalo: Humerus 7141; Ulna
7139; Scaphoid 7128; Magnum 7126; MC2 7130; MC3
7138.

NHML
Dihoplus schleiermacheri
Eppelsheim: Atlas 1284; Scaphoid 1281; MC3 1282.
Dihoplus pikermiensis
Pikermi:  Humerus  M11282b,  11363a,  M48268,  M11367;
Radius  M48168,  M48129,  M11288,  M48253,  M48154;
MC2  M11303a,  M11303b,  M11298a,  M48188,  M48195;
MC3  M11301a,  M11301b,  M48181;  MC4  M11297,
M48187; MT4 M11327.

NHMW
Stephanorhinus jeanvireti
Hajnáčka:  Humerus  1878-46-30;  Radius  1878-46-33;  Pyra-
midal  1878-46-40;  MC2  1878-46-42,  1878-46-42b;  MC3
1878-46-42, 1878-46-42b; MC4 nc.
Dihoplus pikermiensis
Samos  and  Pikermi:  Humerus  1860/0032/0048;  Radius
1863/0001/0025,  1860/0032/0056;  Ulna  1863/0001/0024;
Scaphoid  2009z0089/0001;  MC2  1860/0032/0079,  1860/
0032/0078,  1860/0032/0083b;  MC3  1863/0001/0030,  1860/

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0032/0079, 1860/0032/0082, 1860/0032/0078; MC4 1860/
0032/0078, 1860/0032/0079; MT4 1911/0005/0493.

NMB
“Dihoplus” megarhinus
Montpellier:  Atlas  Mp922;  Humerus  Mp528,  Mp817;
Radius  Mp816,  Mp106,  Mp731,  Mp1032,  Mp329,  Mp453;
Ulna Mp1008; Scaphoid Mp814, Mp324; Pyramidal Mp815;
MC2  Mp985,  Mp526,  Mp646;  MC3  Mp131,  Mp647,
Mp455, Mp525; MC4 Mp733; Pelvis Mp103, Mp333.
Stephanorhinus etruscus
Saint Vallier: MT4 Stv365, Stv243.
Senèze:  Atlas  Se1711;  Humerus  Se1703,  Se1711;  Radius
Se1711,  Se1703;  Ulna  Se1711;  Scaphoid  Se1703,  Se1711;
Pyramidal Se1711; Magnum Se1703, Se1711; MC2 Se1711,

Se1703;  MC3  Se1703,  Se1756  (four  specimens),  Se1711;
MC4  Se1711;  Pelvis  Se1711;  Third  Cuneiform  Se1711;
MT4 Se1703, Se1711.
Upper  Valdarno:  Humerus  Va1680;  Radius  Va1337;  Third
Cuneiform Va612.
Stephanorhinus jeanvireti
Villafranca d’Asti: Humerus Vj89, nc, nc; Radius nc; Pyra-
midal Vj242; MC2 nc; MC3 Vj90, nc; MC4 nc; MT4 nc.
Vialette:  Humerus  nc,  nc,  nc;  Radius  Vt42,  Vt621,  Vt620;
Ulna  Vt621,  Vt42;  Scaphoid  Vt620;  Pyramidal  Vt620;
Magnum  nc;  MC2  Vt621,  Vt621b,  Vt620;  MC3  Vt621,
Vt621b, Vt930; MC4 Vt621, Vt621b, Vt620; Pelvis nc, nc;
Third Cuneiform Vt624, nc; MT4 Vt624, nc.
Perrier-Les  Étouaires:  Humerus  Prr327,  Prr429;  Radius
Prr109, Prr52; Magnum Prr56; MC3 Prr55.