32
HAL Id: halshs-00009680 https://halshs.archives-ouvertes.fr/halshs-00009680 Submitted on 20 Mar 2006 HAL is a multi-disciplinary open access archive for the deposit and dissemination of sci- entific research documents, whether they are pub- lished or not. The documents may come from teaching and research institutions in France or abroad, or from public or private research centers. L’archive ouverte pluridisciplinaire HAL, est destinée au dépôt et à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d’enseignement et de recherche français ou étrangers, des laboratoires publics ou privés. PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM HADAR AND DIKIKA (LOWER AWASH, ETHIOPIA), AND A REVISION OF THE ORIGIN OF MODERN AFRICAN RHINOS Denis Geraads To cite this version: Denis Geraads. PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM HADAR AND DIKIKA (LOWER AWASH, ETHIOPIA), AND A REVISION OF THE ORIGIN OF MODERN AFRICAN RHINOS. Journal of Vertebrate Paleontology, Society of Vertebrate Paleontology, 2005, 25 (2), pp.451- 461. halshs-00009680

PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

  • Upload
    others

  • View
    0

  • Download
    0

Embed Size (px)

Citation preview

Page 1: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

HAL Id: halshs-00009680https://halshs.archives-ouvertes.fr/halshs-00009680

Submitted on 20 Mar 2006

HAL is a multi-disciplinary open accessarchive for the deposit and dissemination of sci-entific research documents, whether they are pub-lished or not. The documents may come fromteaching and research institutions in France orabroad, or from public or private research centers.

L’archive ouverte pluridisciplinaire HAL, estdestinée au dépôt et à la diffusion de documentsscientifiques de niveau recherche, publiés ou non,émanant des établissements d’enseignement et derecherche français ou étrangers, des laboratoirespublics ou privés.

PLIOCENE RHINOCEROTIDAE (MAMMALIA)FROM HADAR AND DIKIKA (LOWER AWASH,

ETHIOPIA), AND A REVISION OF THE ORIGIN OFMODERN AFRICAN RHINOS

Denis Geraads

To cite this version:Denis Geraads. PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM HADAR AND DIKIKA(LOWER AWASH, ETHIOPIA), AND A REVISION OF THE ORIGIN OF MODERN AFRICANRHINOS. Journal of Vertebrate Paleontology, Society of Vertebrate Paleontology, 2005, 25 (2), pp.451-461. �halshs-00009680�

Page 2: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

������������ ���������� ����������������� ����� �!�"�$#&%' �� �(�)���)�� *����+�$�-,$�-,)�.����� /*�

� /0�213�546�����"� �7�"�#849����:�; ��<�81=�>� �?� %@����� ��>A��-�?� %B�C� �� ��.� %D E�>�!���

�������21

���$�F1GA � E�� �H1

I�' �J�KMLON)���� �1P4QL3L)R"S=TEU=T�VXW ��YGZ�R-[\V9�*]3SP^M_=Ta`\43N=b\c�KML$�'� ��81d46%�R-[QeP^aT\49U3fOT6R"[6[\U=gih�ZkjOR�lPmn^6e=R>gom pqR

�S=e3ePZre=fs"Zrs-VXT)taA � ����H1vu= ��-������ ��������$� w%D �� �x�����>� �'�>�

Page 3: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

���$1d�� E��!��� % ]=g giZ�V3R"T��=R>T6gMTMe9s [os�Zkj T6g!]3p=s-_PT!s���])T=sX[Qe3sP�p"R"Z�^a[Qe)R-_dZ eP]3^QTQR ]&g�V�Z ePTa[6fPTagM4��� ������������� ������!

[Qe=U#"$�&%� '����( 49^a]��-]=^6^6S=R Z e2s�_PT ��V�Zr]=^aToePTEU3T��O]=g Z s"g�]3pPs-_PT �$[ U3[QR&%�] R-YH[6s Z�]\eD46�s�_PZX])�PZX[9m��E]9s�_)[6R>]=g T 49ZXe

s-S=R"eP4QpqR>]3Y*�# ��+�,���-�.�� '������ 0/� ��! 1�'2��3�=]3p&s _PT�VX[6s�T �CZX]3^aT6ePT3ma��_=T�p Z�R"g sd]3p s�_dT6giT � VXZX]3^QTMePT�g+�=T\^QZXTagM4

�4 '�+�5�������6 '�����! 7 8�6�������.��/��9%��! ;:=^a[Qe=<OT�U=ZXg s Zre=f9SdZrg8_PTQU2p-R>]3Y s�_PTEVXZkjPZre=f>�4?)(3�� ;�! �4 s ]=�E_DZr^a_HZ s

�&R"]�<&[3<PVrl$f3[6j T R>ZXgMT�Z e$sX_PT�T6[aR>V�ZXTQg sP�7VrT\ZXg s ]3^aT6e=T 4)<3l�Zks g!VXTQgig@�DVr[QfOZX]=V ]��3_P]vU3]3e3sd^6_OT6T'As-T Tos�_Dma� _PTEg8TQ^6]3eOU=4

"$�&%� ����!(CBD�+�� '%��'E): Z g�^\Vr]=g8T\V l$R>TQVX[os�TQU s ]�"$?GFC��%'�G��/���(![aVrs�_P]\S=f3_)YH[Me3l�g��OT9^ ZkYGTMe=gM4\Z eO^\V SPU=Z ePf sX_PT!sXl��PT\4

�$TQR"T��PR-T6jPZX]9SPgMV l$R>TQp"T6RqR"T\U s ]H�� '�+�5�����I�� '�J���! GmQ��_OT s�TQT s _)]3pG"$?KB4�L�� �%��!E$[6R>T�V�Z s�s V TE^6_=[MePf3TaU2R>TQV [Ms"ZrjPT!s-]

s-_O]=g T�]3p'Z s"g��*ZX]=^QT6eOT [Med^6T g s ]3RF4'<=S=sdZ s-g�g�A\SPV�V=UPZrg��PVX[ol g s�_dT [3�O] YH]3RM�3_PZX^�RFT6[aR"R-[Qe=fPT6YHTMe s g s�l!�dZX^Q[QVv] p

"$?�F!�N%����K/��O(o43gFS=f=f=Tag s Zre=f)Zre=^QR"TQ[ g TaU=<=R"]!�$gMZ ePf)g+�PTa^QZ [ VXZ `Q[6s ZX]9eDm ��_=TEg��PVXZ s-<OT sN��TQToe$s-_=T!s���])VXZ e=TQ[afvT9g

�&R"]�<&[3<PVrl�Z eOU=Z ^a[6s TQg�Ta^Q]=VX]3f=ZX^Q[ V3UPZ j T6R"f=TMeP^QT [MePU)^M_P[6R"[Q^Ms-TaR�U=Z�g+�PV [a^QT6Y5T6e3s�<PTos���T6TMeH<PR-]��$g Zre=fjPT6R"g SOg

f=R-[Q`\ZXe=f)g+�PTa^QZ [\VXZr`Q[6s ZX]9edgom

Page 4: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�"���� � �I �!���>� �

�TQfvZre3ePZre=f;�$Zks-_$sX_PT�T6[ R"Vkl$f=T ]&Vr]=f=ZX^6[ V3[6ePU=�=[ VXT6]3e3s ]=VX]3f=ZX^6[ V=g S=R�jdTol�]3p7��[QZXT3<HZ e2s�_OT5K��=Nvc3g

�q��[ Z T3<d4dK��=N9L=#>4\T�vs"TMePgMZ jPT p"ZXT\VXU2R>TQg T6[aR>^o_$_&[ g <PT6TMeH^Q]\e=U3S=^6s�T U)Zre2s�_=T �1�[\g _$jP[QV�V T6l�]3p&�s _PZX])�dZk[E]3jOT6R

s-_OT�V [ g s��vc lPTa[QR"g6m6�"e2s _PT�VX]!�$T6R-�=[ R�s']3p&s _PT4<&[ gMZ eP4 s�_PT!�-e=s-TQR-eP[os�Zr]3e=[ V �p�[9RP T gMT6[QR"^6_2 G�PTaU=Zks�Zr]3eP4\V�TaU <=l

�)m\��])�G�DTMe=gM46�Hm��M]3_P[MePg ]\eD4Q[aePU � m ��[QZXT�<5^6]\eP^6TMe s�R"[os�T6U�]3e2sX_PT eO]3R�s�_OT6Rqe><=[6e�A$]3p&s�_PT��1�[Qg _2 �Z jdT6R'ZXe

s-_OT �$[\U3[aR'[6R>T6[ 4!�_PZ�^M_2lPZXT\VrU=TaU2YH[6e=l$_=]3YGZrePZXU2p-]Pg gMZXVXg 4vZked^aVkSOU=Zre=f s _=T p-[aYG] SPg-��SP^6l&m�

�'[\V�T6]\e s�]PVr]3fPZr^Q[ V R"T\g8T6[QR"^a_HZ e2s _=T ��[ U3[QRD[aRFT6[E^\]9e3s�Zre3S=T\g�s�]=U3[6l;<3ls _PT �$[ U [aR& �TQg TQ[aRF^M_2�'R>] � T6^6s"m6� e

[\U3U=Zrs"Zr]3es ] s-_OT!_P]3YBZneDZrU$UPZ gi^6]3jOT6R>Z�Tag�����]3^'A!��]3]=U)TMs=[\V�mn4\JDcvc3cP4 [aePU2R"TQp-TQR"TQe=^aT\g s�_OT6RFT Zke=#>46s"T6eOg!]3p

s-_O]3S=g [ e=U3g�]3p&j T R�s�T�<PR-[6s�T R>T6YH[\ZrePg�^6]PVXV Ta^6s�TQU2[Ms=�$[\U3[QR��=RF]9j Z UvT�U=[os [�]9e$s-_=T#<PZr]Pg s-R-[as-ZXf3R"[��=_PZX^�[6eOU

�&[ VXT6]vT6e3jPZXRF]\e=YHT6e3s [QV=giTos�s Zre=f$] p=s-_PT�[6R>T6[3mM�C]=g sd]3p&s�_DT�p-]Pg g Z�V&��<DT6[aR>Zke=f�g T U3Z Y5Toe3s�gM49U3[as�TQU s�]$J�m ����� mnb�� [Q4

[QR"T�p"]3R"Y [ V�Vrl$[\g gMZrf=eOT6U s�] s�_=T!�$[\U3[6RP%�]3R"YH[os�ZX]3eP4 �_dZr^6_5^6]\ePg Z�gFs g�]3p&s�_dT p ]&VXV�]!�$Zre=f$YHTQYH<=TaRFg�Z e

[\g ^ ToeOU&Z ePf�g s-R-[as-ZXf3R"[��=_PZ ^]3R-UPT6R t3�E[\g [ VO�CT6YH<OT R �9����#F431dZXU=ZO�$[�AP]3YH[E� 13�$#>4 ��TaePT6e2�$]3R"[E� ��H#>4 ,$[ U3[

�$[\U3[aR�� ,��# m ��_PT6giT YGT6YH<OT R"g�[QR"T!s-_=TQYGg8T\V j T6g�U=Zkj ZXU3TQU)Zre3s�]$e=S=YH<=TQR"TQU)gFS�<���YGT6YH<PTQR"g6m\13s�S=U=ZXT\g�]3p&s _PT

YH[QYHY [ V�Zr[Qe�p�[QS=e=[E]3sX_PT6R&s _=[6eH�=R"ZXYH[os T\g R"TQYH[ Zke5g ^Q[aR"^QT9mv1P]3YHT�]=p=s�_dT �$[ U=[6RDR"_PZ eO]3g �$TaRFT�YHToe3s�ZX]9e=T U

Z e;�=[\g8gMZ e=fH<3l�A�S��QR"ZreH� K��3N��&4&K����\N3#>4 <PS3s=eP]�g lOg s T YH[ s�Z�^�gFs�S=U9l;�[\g�S=ePU3T6R-s-[�A=TaeDm

�"e)[QUPU=Zks ZX]9e$s-]s�_PT!�$[QU=[aRD[QR"TQ[Q4\p"]=gMg Z�V3Rq_PZXe=]2YH[Ms-TaRFZX[QV!�[\g�R"T\^6]3jOT6R>T6U2p"R"]=Y [Qe)[ U�� [ ^6TMe sdgFs�S=U3l

[QR"TQ[E]3e2s�_=T�]����=]vgMZ s T#<=[6e�A$]=p s�_dT �1�[Qg _2 �ZrjPT6R>4\^ [ V�VXTQU��ZOAPZ�A=[\m � _PZXg�[aRFTa[��[ gD<=R>ZXT p Vkl5g SPR�jdTolOTQU)Zre

s-_OT5K��3Nvc=g [ae=Us�_OT6eH<OT fOZre3ePZre=f)Zre K������_=[\g@<=T T6e2sX_PT�s-[aR"f=Tos ]3pD[ s _P]3R"]9SPf\_HgFS&R�jdTol;<3l$[ePT��

Z e3s T6R>U3Z�g ^\Z��dVXZre=[QR�l)R>TQgMT6[aR>^o_;�&R"]��8TQ^Ms"46s-_OT ��Z9AOZ9Av[ �TQg8TM[QR"^Q_2�'R>]��iT6^os�4\V�TaU=< l���TaRFTag TMe&[Ml��V�TaYBgiTQf3T U&ma� _PT

[\VrYG]=g sd^Q]3YH�PVXTos TEg T��9SPTMeP^QT�]3p s _PT ��[QU3[QRP%7]3RqYH[Ms"Z�]9e�Z�g�T��O]3gMTaU2]\e2s-_OT ��Z9AOZ9Av[�gMZXUvTQ4)<vS3sDp Z�TQV�U!�]=R�A

_&[ g�p"]=^oSPgiTQU2]\e2s�_PT 13� ��TaYH<=TQRFm31 ]3YBTEg��OTQ^QZ Y5ToePg �$T6R>T [ V�g8])^Q]=VXVXTQ^os TaU)Zre2s�_PT �gK<=]PVrT�[aRFT6[ 43gi]9S=sX_��

�$T\g sP]=p=�)[ U [aR>49]=p7T6[aRFVrl)YGZXU&U=VXT!�7VrT\ZXg s ]=^6Tae=T [ f=T��-�V�TQYBg8T6fOT6U$[Qe=U)AET6R"[6[\U=gM4QJOc=c3c��vAT6R"[6[ UPg�Tosd[ V�m 43Zre

�&R"T g g #8m

Page 5: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

��_PT �$[ U [aRD[QeOU ��Z9AOZ9A3[�R"_PZ eO]3g�U=T6gi^6RFZ�<PTaU)Zre2s�_DZ gD�P[3�=TQRD[aRFT�_P]3S=giT6U)ZXe2sX_PT!�$[os�Z ]3e=[ V3� SOg T6S=Y

] p=�s-_PZX]��PZ [ 43�U=U=ZXg �1<=['<=[3m\1��OTQ^QZ Y5ToedgD��T6R>T�^a] YH�=[MR>TQU=�$Zrs�_2s�_PT sN�])V�Zkj Zke=f$�p-R>Z�^6[aeHg��=T ^QZXT6gM4

�4 '�+�5�������6 '�����! 7( �9 8�! 0[Qe=U�"$�N%� ����!( F���%'���K/���(a4'�_DZ ^6_)Uv]9S�<3s�VrT\g g�p"]3RqY:[^QVX[QU=T�[QYH]9e=f�V�ZrjPZre=f$Rq_PZre=]=g6m

��_PT p ] R-YGTQR'ZXg�[#�=S=RFT�f3R"[6`vT6R>4'�_PZ�V�TD"$?GF!��%����+/-��( Z�g�YH]vg s"V l$[#<&R"]!��g T6R7p-T\TaU=Zke=f$]3e5V TQ[MjOT9g�[MePU sN�$Z fOg6m

��_PT �$[ U [aRD[QeOU ��Z9AOZ9Av[ R"_PZreP]3g�_&[MjOT [\VXgi] <PTQTMeH^Q]\YH�3[ R"T9U;�$Z s�_Hp-]Pg8gMZXV�g�p-R"] Y:[�ePS3YH<PTQR']3pdV�]=^Q[ V�Z s"ZXTagM4

s-_OT YH]Pg sdZXYH�=]=R�s-[ae3sd]3p �_dZX^6_)[aRFT=t

��[6s T ��ZX]v^QToePTvto�7Z�A=TQR-YGZ�431P[6YG]=gM46�7T6e3s [QVX])�v_P]=gM4Q��_=T\g8g [ V�]9eOZ9AOZ&�-A�R>TQT\^6T�� �R-[QY$<P]9SPR-f2[QeOU

�7Zkj T6s�TQ[MSD4&K��3J ���vAT6R"[6[ U&gi4 K�� �����\ATQR-[Q[ U=g�[Qe=U ,�]3S=p ]=g 4DK����=c3# �Q,�T6YGZ9AOV�Z s T��PT���� S=RMA=TMl �vAT6R"[6[ U=gM4

K����\L&# �a��] S=[aRFZX[E�-��S3ePZXg ZX[ �vA�S��aRFZreP4&K������ # �a��ZXgMg [ g p-[��� ]3RF]v^Q^a]��\S=e!�=S!<PV�ZXg _PTaU3# �6��[ Rq[af=_PTo_5���"Rq[ae �

��_PToePZrSOgM4dK��=b3b��\AT6R"[a[\U=g 4DK������\#8m

��VXZ�]3^QTMe=T=to,�]=]!<PZ9%�]3R-[\46/.TQg s ��S&R�A=[ae=[\46,$[ae&[��P]=ZX4 ��]9s�_=[Qfv[6Y ��,�T6e3lO[ �Q��]=]=Z �8TQRD[QePU �'[Ms�s T6R>g8]3eP4

K��3N\J �a�$[6RFR"Z�gM4&K��=N��=4 K�� �����a�$[9R-R>ZXg�[aePU ��TQ[�APTolO49JPc3c��\#��\��YG] <$[ V�V�Tol5� �s�_dZr]!�PZ [ �6��]P]=Z �8TQR"4DK��3N����

��]=]=Z �8TQRD[aePU)��_vS=RF^M_PT6R>4&K����3b��vA�S��6R>ZreP4dK����3b\# �\��_dV3[QVd��S=f3_PVr[QY ���*]3R>]3^a^Q]�� S&eG�3S�<=V�ZXg _=T U3#

��VXT\Zrg s�]=^aTMeDT3t ��SPV [ U �$[aYBZ�U3[E� �C]3R"]3^Q^Q]�� ATQR-[Q[QUPg�[Me=U ��T6R"e=]\SPg g8Z�4\Z e>�=RFTagMg # �\�V�U3S3jP[QZ�4 ��T6ePZre��

�q��[MeP`Q[6ePZr[��a��TQ[�AOTolO4 K����&b��a��]&]=Z �iTQR"4&K������ �\ATQR-[Q[ U=gM4 K����\N3#8m

��_PT YH[os T6R>Zr[\V=pqR"]=Y+��[\TMs-]=V�Z3[6eOU ��[ae&fvT�<=[a[6e���T6fP4'�3S�<OV�ZXg _OTQU=<3l�A�S��QR"ZreH� K�� �vN3#D[aedU �$]v]=Z �iT6R

� K��vN3J=#F4QR>TQg��PTa^6s ZkjOT\V l 4'�[Qg�eP]9s U&ZrR>T6^6s Vkl�T3P[6YGZ eOTQU&mM� T6[\gFS&R"TQYHTMe3s g [aR>T�Z e�YBZXV�V�ZrYHT6s�TQR"gM4Q[6ePU)p-]PVXVX]!�

A�S��6R>Zke5� K�� �vc3#8mo��ZX[6fveD]vg TQg�V�ZXgFsd]9edV l�UPT6R>Z jPTaU2p"T6[Ms-S=R>TQg6m

������� �������������������������������� �����3� !#"%$'& 4 �$[Ms-S=R"[QV9��Z�g s�]3R�l� SPg TMS=YB4Q��]9eOU=]9e �)(+*�,E4 % [Q^6SPVrs �

U=TQg!1=^\ZrT6eO^\T6gi4 ��lO] e ���-$.*��'/�49�-ePg s�Z s�S3s=�$[as"Zr]3e=[ V&UvT6g!1d^QZXToeO^QTQg�UPT�VXW �RF^a_��6]=VX]3fPZrTET s U9S2�7[Ms-R"ZXYB]=ZreOTQ4

E['<=[os �10.$'".46�$[6s�Z ]\e&[ V3�wSDg8T6S=Y5g�]3p&,�Tae3l=[ 4 ��[\ZXR"]!<DZ �)"324"5,E4 �wSdg �QT�A��a]=VX]3fPZ �3S=TET s

�*Zre��6R"[QVX]3fPZ �3SOTQ4 ��[aSPg [ e3e=T���"6$'&.$'/�4 � S=g �oS=Y �$[6s Z�]9e=[\VvU&W ��Z�g s�]=Z R>T �$[Ms�S=R>TQV�VrT\4 � [QR"Z�g��)"678��4 � [QU=Tae

��T6s&APZ A jdT���R"[QYH[ � SPg8T6S=YG49��e�Av[QR-[��1$'&9!�46�$[6s�SPR-_DZXg s�]=R"Z�gi^6_=Tag �wSdg8T6S=Y �E[ g TQV��:$'&9"%; 4

�$[6s�S&R-_PZXg s�]=R"Z�g8^6_OTQg �wSPg8T6S=Y / Z�Toe �:$<"3=�46�$[6s Z�]9e=[\V9��SPg TMSPY ]=p&�s _PZ ]��PZ [ 49��U=U=Z g�� <=[�<=[��:$<">7E4

Page 6: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�$[6s Z�]\e&[ V9��SPg TMS&Y:]=pD��[6e=` [6ePZX[Q46�$[QR � �gJ� 1O[\Vr[Q[6Y �:/ ���<".46�'[ V�� ]9e=s�]&V ]3fPZrgM^o_PTag��"ePg s-Zrs�S3s-4Q���ve=g s�TQR �

*�"6$.*P4=13s-[a[6s V�ZX^6_OTQg�� SPg8T6S=Y p��&RP�)[Ms-S=RMA3SvePU3T\4313s-S3s�sXf=[6R-s"m

� ��1� �������1����� ���� �����J� �<,E4\�p-[9RP��]=^ [ V�Z s�l�� �$[ U3[QR'g [QY$�PVXT6# ���<��0G4 ��Z�APZ�A=[���=� �4 ,�]=]!<OZ9% ]3R-[��

; 7E46/.T g sP��SPR�A=[6e=[3m

1��H1=���� ����>� �'� ��� �� ���������A �

%'[ YBZXVrl �$�-������ ������-�)���A R"[6l 4 K �QJ�K

1OS�<=p-[6YGZ�V l$ ��2�-������ ���� ���)���ARq[6lO4�K � J�K

��R"Z�<=T �$�>��� ������-�2�P �Z e=fPg s R��=YG4dK��3J3LP4vT6YGToePU&m

ATMe3Sdg ���������������������! #" A�R-[ lO4DK�� � �

$ '�K �/&%- �9��%' ����!(���Z�TMs-R"ZX^o_d4&K��3LvJ

7('*) � �+) �-, � �1�J� �� '������������ ��3���� (��� 8�! :�9�ESPR-^6_PTaV�V�4dK��DKaN3#>4\V�Z jdZre&f$�p-RFZ�^6[ae/.9�_PZrs"T10vRq_PZre=]&m

��� �12�������� �3� �$[\g [ V3[6ePU2p-R>]3e3s�[\V3_=]3R"e=g6mM�$[\g8[\V!<=]3ePTag�RF]\S=e=U3T U2[6eOU)g _O] R�s�4Qe=]3sd^6]3e3s [a^ s Zre=fs�_PT

V [ ^QR"Z YH[QV-mM��]C�$T6R�<P]3R"U=T6R7]3pD]3RM<PZksdgMVr]!�OTQg�U3]!��e��[aRFU=gMmo/0T6[�A=�d]3gFsM�"]3R�<PZ s-[\V!�=R"]v^QTQg8g6m3��R>]3[\Ue3SO^6_=[QV

^QR"TQg s>mo� RFT6YH[� Z�VXV [�YHSP^6_)R"T U9S=^ T6U&maI;�!�=TQRD[ ePU)Vr]��T6R'Z�eO^\Z gi]3RFg�jOT\g s"ZrfOZk[ V=]3R�[�<Pg TMe=sqma�'[aRq[ ^\]9e=T�p"]=V�U

�$TQ[�APm\��e3s"Ta^QR-]=^6_OT6sP[3<Dg8T6e3s"m

�� ��2����-V [Ms"T!�*Zr]=^aT6eOTQ4 �[ g s T R�e2� TQU=Z s T R-R"[aePT [Me �\Vr[6s�TQg s=� Z�]3^QTMePT s ] ET ^6T6e3s�4Q�pqR"Z�^6[3m

��_PT fvT6e3SPg��4 ��+�,�����I�� '�3���! 54Q[Qg�_PT6R>T SvePU=T6R>g s ]3]=U=4\Z�g@�v[QR-['�=_3l V T6s ZX^Q49[aePU)Zre=^\VrS=UvTQg�[QV�V9��Z�^QTQR-]9s�ZrePZ

T�=^ T��9s�"$�N%� ��K�!(a4Qs-];�_PZ ^a_HZ sPf=[ojdT�R"Z�g TEZre2s�_PT Ta[QR"Vkl�7V-Z�]3^aT6e=T\mQ��s"g�T6[QR"V�Z�TQg sPR"T'�=R"TQgiToe3s [os�ZkjOT�Z�g

�4?)/� ��C 1�32��3�=pqR"]=Y+s _=T�VX[os�T �CZ ]v^6Tae=T�]3pPs�_dT �[ g s�TQR-e�*TQU=Z s TQR-Rq[ae=TQ[Me'mQ��eP]9sk_DT6R ]!�3s"Zr]3eP4QeP] sdp"]=V�V ]C�$TaU

_PT6R>TQ4'�$]3SPV U8<PT s ]2R"T\gFs-R"ZX^osPs�_dZrg�fPToeOTQR"Z�^!e=[QYHT!s-]��VXZX]3^ ToePT�[MeOU � V T\Zrg s ]=^6TMePT�p-]=R-YGgM4 [\pXs"T6RDs-_PT "$��%' '�K�!(

g��PV�Z s�4Q[Qe=U s�]=AvTQT��>�4? /� ��! ��2����P[Qg�[#�DVXTQgiZ ]\eD4\]3RDs ])TQR-TQ^MsP[ edT�� fvT6ePTaRFZr^�e&[aYHT�p"]3R'Zks>4Q[ g�eP]9ePTEZ g

[6j=[ Z�VX[�<PVXTE�qAT6R"[a[QUPg 4DK�� ���Pt J=L=# m

Page 7: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�����*��������� ������ #" " � ����+����� �J� *" ���7]9YGT\VX4 K������\#

�4����/���%� '�K�C(�� �H� �� �����!( 8�6�������.��/��9%��!(!� ]3YHTQV�4dK ���3bPmM��]3YHT6e)evSPU9S=Y m

�4����/���%� '�K�C(� �6���3���I��/6�&%��)(��7]3YHTQV�4dK �����dt J�Kac&m

$ '�K �/&%- �9��%' ����!( �� �)��%�� E ��ZXTos R"Z�^M_D4&K��3LvJDm

, �!,:��� '�) ��� �']=YGT6V=� K �����\#7U=ZXUe=]9sdU=TagMZkfve=[os T�[ s�lG�DTQ4QeP]3R'U=ZXU2_OTVXZ gFs&s�_PTE^6]\e3s-Tae3s-g�]3pPs-_PT s�lG�PT'�

gMT6R>ZrT\g p-R>]3Y s-_OT �7VXTQZ�g s ]3^QTMe=T]3p=�'[\VXZ�A3[\])���@��TQR-edZrp"Z ePTQ4Qe=]!�0��ZXf9_OT6ePZ p 43�VrfPT6R>Zr[9#"4!<=S3sPsX_PT6R>T�ZXg�eP]2R"TQ[ g ]\e

s"]8<PTQVXZ�Toj T sX_P[Ms=s-_=TEg��OTQ^QZ Y5ToeOg s _=[6s=_PTZXV�VnSPg s-R-[as-T U)Zre K � �&b8�$T6R>T eO]9s��=[6R-sd]3p7Z sFmQ�7_PT6R>T<=l�g TQVXTa^6sP[\g

V�Ta^6s�]OsXl6�=T s _=T!S!���OT6RD��JH� ������ eO] m ��� ��J3J �DKa#�p>Z fvS=RFTaU$]3e)_PZXg �7VX[os�T2K 4 % Zrf=S=R"TQg K��-JDm

��� �12�������� �3�q1dZ `\T�Vr[QR-fPTaR&s-_=[Qe5Z e$�4?)/6 ��� ��2-�J�qmM�$S=^a_=[\V3^6R>TQg sdgFs-R>T s�^o_OTQU2YH]3R>T�^6[aS=U3[ V VrlPm6�$[ g [QV

eP]9s ^o_HgF_P[QV�Vr]��T6R8mM�'RFTaYB]PVr[QRDR"]�� g _O] R�s�ToeOTQU&m\��R-[QeOg jPTMR>g T�V�])�=_Og�]3pDS��G�PT6R&s�T6T6sX_HV�]9ePf2[MeOU2eP[6R"R"]C�)m

�*Tos [QVX]��=_)Tvs T6e=U=g�UPZXg s�]=VXZke=fvS=[QV�Vkl�Zre=s-]s�_OT�U=ZXg s�[\V=^QZ evf=SPVrS=YB4\^\Vr]=giZke=fs-_OT4�P]=g s-p"]=g8giTos�s T9m

�� ��2����-V [Ms"T!�7V�Z ]=^6T6ePT!s ])Vr[6s T � VrT\Zrg s�]=^QTae=TQ4Q��]3R�sX_��pqR"Z-^6[��QYGZ�U=UvV�T ��V�ZX]3^aT6e=T!s"]$T6[QR"Vrl

�7V�T Z g s ]=^6TMeDT 4 �[ g sP�p-RFZ�^6[ �vVr[6s T!��VXZ�]=^QToeOT\4O1 ]3S3s�_��p-RFZ�^6[9m

��� � ,)� � ) �� ���9���� �8, ����� � ��� ��� "5����)�:� ���N�q1�A=SPV�V9�2�-, � K�� Koc)�F13� KQ#it s-_PZ�g�g�A\SPV�V!�[ g�p-]=S3eOU

[\g gi]3^QZX[os TaU=�$Zks _)[ �8S jOTMeDZXVXT _O]3YBZneDZrU�g�A3T\VrT6s�]3eDma�-sdZ�g�p-R ]3Y [6e�]&V U)Zre=UPZ jdZ U3S=[QV��6s-_PT�Y)SP^M_H�$]3R-e2��JPg [ e=U

� �=gD<PTaZre=f s _=T�]3eOVkl�R"TQY)[\Z ePZne&fs�TQTos�_Dm ��_=T YGTQUvZrp"]=gMg T6sXs TQg�[QR"T^QVX]3g8TQU&m ��_=T YGTos [QVX]��=_)]3p=��JHZ�g

e&[aR"R"]!�$49[aeDU$gMVXZkfv_3s"V l�]�<PV Z��\SPTQ4)<vS3sdZks g [3dZXg�Z�g YG]3R"T#�=]=g s�T6R>Zr] RPs-_=[Me2s _=T Y5TQU=Z p ]&gig Tos�s�T3mo� �)ZXg

s-R-['�PTa`Q]=ZXU3[\VX4)�$Z s _)[#�$Z�U3TEToe=s�Rq[aeP^aT s ] s�_PT YGTaU=Z gMZne=S=gom ��_=TEV [�<DZr[\V!�[\VXV=Z�g�^Q]\e=^Q[Mj=T <OTo_dZ ePU2[�R"T\U9SO^QTaU

�&[aR"[Q^a]3e=T�p"]=VXU&m ��_PT��=[QR ]=^Q^QZ��PZks�[\V��=RF]v^QTQgig T\g�[6R>T�VX]3e=f[aePU)g VXTMePU3T6R8m � _PT4<=[\giZX]3^a^9Z��PZ s-[ VPZ g <3S3s g V�Zrf3_3s"V l

[Qe=f=VXTaU2R>TQV [Ms"ZrjPT s ] sk_DT4<=[ gMZ g+�v_PToeO]&ZrUDm ��_PT!e3SP^6_=[ V=^6R>T6g sdUv]=T6g�e=]3sdT3s T6eOU2^Q[MSOU [ V�V l$YHSD^M_H<OT lO]\ePU sX_PT

]=^6^ ZO�PZrs-[ V=^Q]9eOUOlOV�T6g6m

1 AvSPV�V3��� � K J�����JDb$�q1=� Ka#'Z�g�s-_OT YH]=g sd^a]3YH�PVXTos"T�gKA3SPV�Vv� %�ZkfDmPKa#F4QYGZ�gigMZre&f�]9eOVkls-_PT�RFTaf=ZX]9e

[Qe3s T6R>Z ]3RDs ] � �$[6eOU2YH]Pg sP]=p=s-_PT e=[ g [ V!<=]3e=T g6mo�-s g�YG]3RM�=_=]=VX]3f3l�ZXg �3SPZrs�T�gMZXYHZ�VX[6RPs"] s�_=[Ms']3p=s-_PT�YH]=U=T6R-e

p"]3RqY@m3�]9s�_H�$ZXe=fvg!]3p s _PT evSP^o_=[\Vv^QR"T9gFsdg s�R"Tos ^o_H^Q[MSPU3[ VXVkl;<PTol ]3e=U s�_=TE]=^6^\Z��dZ s-[ V=^6]\ePU9lOV�TQgM4'�$Zks�_)[EU=TaT��

Page 8: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

g [6fOZks�s�[ V3e=]3s�^6_Dm �!_=TQT'As�TQTos�_)[aRFTEZre)YGTQU=ZrS=Y*�T6[QRFm ��_OT�VX[�<PZX[QV��[QV�V=Z�g�g Zre3SP]\S=gM4'<=S3s=s-_OT6R>T�Z�g�eO]

�&[aR"[Q^a]3e=T�p"]=VXU&mQ� V�]3e=f�^QR-]=^M_PTosdZ�g �=R>T6giToe3s ]9e)[ V�V9s�T6T6sX_d4Q[Qg��$TQV�V3[ g [1�$TQV�V��"U3T6jPT V ]��PT6U$^6R>Z�gFs-[E]9e2s�_PT

YG]=V [QR"gM4\TMeP^QVX]=g Zre=f s�_OT�YHTQU=Z p ]&g8giTos�s�TE]3e2sX_PTEV TQpXsDYG]=V [aR>gom ��_=T4�=R>]9s ]=V ]��3_HZ�g�VX]\e&f$[6eOU eP[aRqR"]!�Hm ��_=T

�P]3gFs"T6R>ZX]3R'^\Z e=f3SPVrS=Y ZXg _PZrfv_P4 [ e=U s _=T��P]vg s p"]=g gMT s�s�T�Zrg�^\Vr]Pg T U2]\e)[QV�V9s"TaT6sX_HT�O^QT�3sP� �P4'�_PZ ^a_2_P[Qg�[

s-R-['�PTa`Q]=ZXU3[\V=]\S3s-V�Zre=T9m

1 AvSPV�V3��� � K JDbG� KE� 1v� KQ#'Z�g [ V�gi]2[ae)[QVXYG]3g s ^a]3YH�PVXT s T g�A\SdVXV�m ��_PT�]9ePVkl�gMZrf=ePZrp"Z�^a[Qe3sPU=ZXpqp T R"TQeP^aT

pqR"]=Y+s _PT4�&R"T6jPZX]\S=g�]3e=T�Z�g s�_=[Ms&s�_PT�]=^a^9Z �DZ s-[ V3e=]9s ^M_HZ�g�g8_=[QV�VX]!�)mQ��_=TEU3T��3s�_�]3p s _PZ�g�e=]3s�^6_)_=[ag�e=]

g��PTa^\Zkp"Z�^ jO[QVXS=T\4Q[\g�g8_P]!��e;<3ls-_PT!jP[6R>Zr[6s Z�]9e5Z e2sX_PTEV�Z jPZre=f$p"]3RqY �-gMTQT!��]=]3g8T\4&K��=N=bPt �PV�maJP#Fmv��_=T s�T6T6sX_�[QR"T

YG]3RFT#�]3R"eP4)<3S3s gF_=[QR"T�[ V�V9s-_OT p"T6[Ms-S=R>TQg�]=pD� � � KaJ�����JDb3m

��[3dZ V�V [�� � � K��3N���J�� � 13� K #'ZXg!p�R>]3Y [ lO]3S=e=f�g��=T ^9ZnYHT6eP4'�$Zks _2�DL � S=gFs'TaR-S!�3s T6UDm6�-s g _P]!�$g�s�_=[6s

s-_OT YGTQU=Z p ]=g g8T6sXs-TE^\Vr]=g8TQg!T6[aRFVrl�ZreH�$TQ[6R>46s-_=[Ms&s�_dT6RFTEZXg�eP] �=[aRq[ ^\]9ePT�p-]=V�U�TojPTMeHZre2s-_OT!S!���=TaR �=[aR-sd]3p s�_dT

^QR"]!�eP4 [ e=U s�_=[Ms&s�_PT s�R"[6edg jOTQR"g TEV�]��3_Pg�[6R>T�VX]3e=f$RFTQVX[os�Z jdT s ] s�_OT9ZrRDs�_DZ ^ A\eOT9g8g6mM��!KEZ�g�e=]3s��=RFTag TaR�j T6UP4'<3S=s

�[\g�^QTQR�s-[ ZkePVrl;�=RFT g TMe3s'Z e5V�Z p T9m

��]=g s ] p=s�_DT�YGT6[\gFS=RFTMYGT6e3s g�]3p&s-_PT g TEg�A\SPV�V�g�����[3<PV T)K 46%7Z fPm6JP#'p-[\VXV �$Z s-_PZre2s _=T R"[ae=f=T�]3pPYG]&UvT6R"e

�4?)(3�� ;�! Hma��_PT YH[QZreHT3P^aT��3s"Zr]3e5Z g�s�_=T�e=S=^M_&[ V=^QR"TQg s�4 �_dZr^6_5Z g <PR-]=[aU=T R Z e2sX_PT!�$[QU3[ R'g��=Ta^\ZkY5T6e=g

��YGT6[ g S=R>T6YHTae3sPeP]&m K b\# m

� ��� ��� � �$�-YGT6YH<PT6RdS3e�A9eO]C�Ee=#'Z�g�[�p"R"[Qf3YGT6e=sP]3pD[EYH[�PZ�V�Vr[#�$Zks-_$��'J�� ��dLDm\��_OT�T�\s"T6R-e=[ V��[QV�V g

] p=�� ������'L)[QR"T�fPT6e3s"Vrl$S3eOU3SPV [Ms-TaU=4)<vS3s&sk_DT4�=[aRq[ ^Q]9eOT�p-]=V�U)Z�g �$T6['APmQ��_=T�^6R>]3^a_=Tos�4 [ R"Z�g ZXe=f)p-R>]3Y s�_PT

YGT s�[\V ]!�3_H^QVX]vg T s ])Zrs g �iS3e=^6s ZX]9eH�$ZrsX_2s�_PT�T ^ s ]=VX])�=_P4QYGTQTos�g sX_PTE^6R>Z g s [!s�])T6eO^QVX]=g T [�R>]3S3ePUvT6U

YGTQU=Z p ]=gMg T6srs�T�]3p'g YH[\VXVPg Z ` T9m��E]9s�_2sk_DT4�=R>]9s-]=VX]��3_)[ae=U2YGTos [QVX]��=_)[6R>T�^MSPR�jPTQU=4\[6eOU s-_OT YHT6s�[\Vr]!�v_

^Q]9e3s�Zre3S=T\g@�O]=g s"TaR"Z ]3R>Vkl�Zre=s ]2[�_DZrf3_5U3Z�gFs-[ V=^QZ ePf\SPVkS=YG4�<3S3sPsX_PT�VX]!�3_Og [ R"Tg _=]=R�s T6RDs�_=[QeHZ e$�#?'(3�9 �! Hm

��_PT6R>T�Zrg�[Eg8Y [ V�V!�P]=g s�p"]=gig TMs�s T�]\e2��DJ'm

">������� ��� �)� ����� , ���<�1�+7 �1�:-�"�VXV&VX]��$T6R&s�TQT6sX_)p-R>]3Y+s-_OT �$[QU3[ R&%7]=R-Y [Ms"Z�]9eHVX[Q^�A s�_=TUvT6R>Z jdTaU

p"T6[6s�SPR"T9g�s�_&[MsP^a_=[6R"[Q^6s TQR"Zr`\T sX_=]=giT�] p= �TQ^QTMe s��4?'(��� ;�) �4\g SP^o_�[\g�ZreO^6RFTa[\g8TQU2_3lG�Pg8]3U=]9e3s�lO43R"T9^Ms-[Me=f3SPVX[6R

Page 9: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

]3S3s V�Zre=TQ4\[6ePU�^QVX]3g8TQU2p ]&gig Tos�s-ZXU=g6m\1PZre=^\TEg ]3YGT�]3p=s-_PT\g8T s"TaT6sX_)[QR"TE^\T6R�s�[\ZneDV l�]=p5�4? 8�����K�����!/���%��C �46s _PTol

g _O]!�;s-_=[Ms&sk_DTag T�p-TQ[os S=RFT g�_=[\U eO]9s&lOT6sdToj ]=VkjOT6U�Z e2s�_PZXg�VXZ e=TQ[af=T3mM��]C��TojOTQR"4 s�_DZXg�[\VXg ]=�=R>TojOT6e3s g

U&Zrp"p-TQR"TQe3s"Zr[6s ZX]9e;<DTMs��TaTMe2s�_PT YH[6eOU&Z <PV T\g�]3p=s-_PT!s��$] s [�=[#�&R"T\g8T6e3sdZre2s�_PT �$[ U=[6R&% ]3R-YH[6s ZX]9e'm � s-_PT6R

p"T6[6s�SPR"T9gi4\g S=^6_)[ g�^Q]3R&�=SPg�U=T3�3s�_H]3R'T�3s TMe sd]3p=s-_PTEg l3Y$�=_3lOgMZXgM4Q_=[6jOT eP]9s��PR"]9jPTaUSDg8T6p-SPV&p ] R g��PTQ^QZXp"ZX^

U&Z g s-ZreO^ s ZX]9ed4'<3S=s=s�_PT ^a]3e=U9l VXT�]=pD� � � KaJ&b6� KEZ�g@�=[ R-[\VXV�TQV9s ] s _=T�]&^a^QVrS=g [ V��PV [Me=T�[\g�Z e>��?'(��� ;�� �4 �_OTQR"T [ g

ZksdgMVX]!�=Tag�YBT U=Z ]���jPTMe3s�Rq[ V�VrlHZre "$�N%� ��K�!(�F!�9%��!�K/��9( mQ�V s _=]3S=f\_)[#<DZ ] YHTa^M_&[MedZX^ [ V3[ae=[QVrl=gMZXg �[ g�eP]9s

S=e=UvT R�s [�APTMeP4 s�_dZrg�] R"Z�Toe3s�[6s ZX]9ed46s"]3fvT6sX_DTaR-�$Zks _ s�_=Tg8YH[QVXV�TaR'gMZr`\T�]3p&s-_PT4�P]=g s �-f=V�ToeO]&ZrU;�&R ]=^QT6gMg�[ae=U

pFVr[6s�edT6gMg�]3p=s-_PTE]=^6^\V SPg [ VPg S=R"p-[ ^QT\g 49[aR>T�^QV�T6[QR"Vkl�VXZXe�APT6U�s"] s�R"[6edg jOTQR"g TE^o_PT��$Zke=f$YH]9j T6YGT6e9s g!� %7]=R�s"TaV�ZrSPgi4

K�� �vb3#Fm

/ ��� ,1�:����� �����J�-� [3P]\e=]=YBZ ^�UPZrg s"Z eP^Ms"Zr]3eHZXg�[\VXg ] jOTQR�l5U=Z p"p"Z�^6SdV sd] R'ZXY$�=]3gig Z�<PVXT�p-]=R'V�ZrY$<H<P]\e=T\gom

��_PT4�=RF]��=]=R�s"ZX]9ePg�]=p s _PT�^6R"[aeDZr[\V3YH[Ms"T6R>Z [ V=] p�<P]9s�_2s-[3v[g8S=f=f3T g s=s�_=[asPYB]vg s��O]&gFs-^aRq[aePZ [ VXg [QR"TE] p

�4 '�+���O�����6 '�����! �4)<vS3sDeO]2f3R>T6[6sdgiZr`\T�U=ZXp-p"T6RFTae=^QTEg _P]\SPVXU=<=TET�G�OTQ^os T6UDm ��ePVkls _PT U=Z�g s [QV=R-[\U=Z�Z=gF_P]!� [

U&Zrp"p-TQR"TQe=^QT�]=p �=R>]!�P]3R�s"ZX]\ePgM4 s _=T!s��]�g YH[\VXVXT6R7]9ePTQg���"$�&%� ����!(��M# <PT Zke=f$R>TQV [Ms"Z jdTaVkl;�$Z�U=TaR s�_=[ae2s�_PT�VX[6R"f=T6R

]3e=TQg6mo��_PT�<=Tag sdg��PTa^\Z YGT6eHZXg�[Me)[ gMg8]=^QZ [Ms T6U�g Tosd]=pDV�ZrY <;<P]9ePT g 4Q� ����� � J�� b � ��$J&#Fm � _OT _=S=Y5T6RqSOg [ e=U

p"T6YHS=R'[QRFTE^QVX]=g TEZ eHV�TMePf9s�_2s�] s _=T�Y [�PZ Y�S=Y R"T9^a]3R"U3T U2p-]3R7VXZkj Z e=f>��?�(3�� ;�! �4)<3S3s=s _=T�Rq[ U&ZnSPg 49SPVre=[ 4 s Z9<PZX[Q4

[Qe=U$YBTMs-[3�P]3U=ZX[QVXg�[QR"T�V�]9e=fOT6RF4\g _O]!�$Zke=fs-_=[as=s�_OT�U=ZXg s [QV=VXZXY <5TQVXT6YHT6e3s g �$T6RFT�YH]3R>T�TQVX]9ePf3[Ms T6UP4Q[Qe=U

Z eOU=Z ^Q[Ms"Z ePf$[ YG]3R"T^MS=RFg8]=R"Z [ V YG]vU3T�] pdVX]&^a]=Y)]3s Z ]\eH�"��['<PVrT�J=#Fm

�# ��+�,���-�.�� ��3���� 0(��O �C 0�&��S=R>^M_PTQV�VX4 K��&K N3#

�"e)[QUPU=Zks ZX]9e$s-]s�_PT!sN�])V�ZkjPZXe=f)gFS�<Og��=Ta^\ZrT\g 4Qs�_PT�p-]Pg gMZXV3p>]3RqY �4�6�O/6��%' ����!(�(��O �!(

%- '�� 1��/���� �'�3��%3�!/��!(���Z�V ` _=T\ZrYHTQR"4dK �9JPb=4\Z�g�ZXe=^QVrS=U=TaU&m

��� �12�������� �3�q1=s�R>]9e=f8�P]=gFs�]=R�<PZks-[ V=^Q]\ePg s-R"Z�^os Zr]3eDmM�)SP^M_P[QVv^ R"T g sPeP[6R"R"]C�)mM�7]Pg s fvV�TMe=]PZrU �PR"]=^aTQg g

�$TQ[�APm\�!R-[QePZ�]\YH[6ePUvZO<vSPV [aR'[QR�s Z�^6SPVr[6s Z ]\e)_P]3R>Zk` ]9e9s [QV-mMIH���OT6R7^6_=TQT�A s TQT6sX_$_3l��Og ]3Uv]3e3s"4'�$Zks�_2jdTaR�l�VX]=e=f

e&[aR"R"]!�.V�]��3_Pg�[6ePU)TMePV [aR"f=TQU2p ]&gig Tos�s�TQgMmo��[�<PZ [ VC�[ V�VXg!g8Z evS=]3SOg�ZXe2sX_PT SG�!�PT6R��&[aR�sd]3pPs�_dT�^aRF]!�EeDm\�^a^QVrSPg [QV

Page 10: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

g S=RFp�[\^QT�p"V [6s"m6�7R"TQYG]=V [aRDR>]!�.g _P]3R-s T6e=TaU&46��!K g _=T U <OT6pF]3R>T [ U3S=VrsX_P]3]vU&46�'J5g Y [ V�V�m ��e)YH]=V [aR>g 4\^QS=R�j T6U

�&R"]9s ]vV ]��=_P4\]!<PV�Z �3SOT e=[QR-R>]!� YGTos-[ VX])�=_P4\^\Vr]=g T U2YHTaU=Z p"]=g giTos�s T94'�O]&gqsd[6eOU=�&R TQp-]Pg g8T6s s T\g�^QVX]=g TaU)Zre

[\U9j=[aeP^aTQU=�$T6[QRFma��]!�TQR'^6_PTaT�As TQT6sX_)R>T6^6s�[6e=f\SdVr[QR"4)�$Z s"_H^QVX]vg TaU2p"]=gMg8T sXs�ZrU&g�]\eH�$]=R�e2s�T6T6sX_'mM�*T s-[��P]3U=Z [ VXg

g _O]3R�sP[QePU�g s ]9S3s"m

�� ��2���� ��]��$T6R&� VXTQZXg s�]=^aT6e=T!s"] T6^QTMe3s"46�[\gFsD�p-R>ZX^Q[ �vYGZ�U3UPVrT!��V T9ZXgFs�]&^aT6eOT s ] ETQ^ Toe3s�4O1 ]9S3s�_

�pqR"Z�^6[��\VX[os�T �7V�TQZ�gFs-]v^QTMe=T\46��]3R�s _)�p-RFZX^Q[\m

�� ���)� �:� � * )��!,1��� �:���3� %7]=g8gMZ�V3R"TQYH[ ZkePg�]=p��� ������������� ��K�&�! .p-R>]3Y s�_PT �7V�TQZXg s ]3^QTMe=TE] p=s-_PT!��]!�$T6R

� �[\g8_2jP[ V�VXT6l$[QRFT!s ]=]5p�Rq[af3YGTQe3s-[aR-l2s ]=<PTEZ U=TMe3s"ZXp-Z�T6U s�])g+�PT6^\ZrT\gi4 <=S3sP[ R"T��PR-]!<=[�<PV l$[ g gMZrf=e=[3<DV T s"]

�4?)(3�� ;�! Hma�Cs"[ V SPg�pqR"]=Y ��g�<P]&V T94\g8]3S3s�_��TQg s ] p=��[QU=[6R>43Z�g�Vr[QR-fPTaR��-YH[dZrYHS=Y*<=R>T6[QU9s�_ � KvKac3YHY �

V [Ms"T6R"[ V _PTaZ f\_=s � � �QYHYH#Ds�_&[ae2s�_=T�Y [3dZrY)SPY R>TQ^Q]3R"U=TaU <=l�A�S��6R>Z eH� K����\c3#�p ] RPs�_dT�VXZrjPZre=f2p"]3R"Y mQ�-s

^Q]9e=p"ZrR"YGg s _PT�gMZ `\T�R"T\U9S=^6s ZX]9e�] p=s-_PZ�g�gK�PT6^ Z T g�giZkeP^aT s�_OT YGZXU3UPV T!� VrT\Zrg s�]=^QT6eOT�����]3]=Z � TaR"4 K������3#8m

� ���/)����:� �-�����

�� ��L�,�����I�� ������! 7 8�����J�����!/���%��C �46s _=T�p"ZrRFgFs'p-]=gMgiZ�V3R"_PZre=])T6jOT6R'e=[6YGTQU$p-RF]3Y �p-R>Z�^6[ 4!�[ g�UPT6gi^QR"ZO<OTQU

]3e)p-R"[6fvYBTQe3s [6R-l)R"TQYH[ Z ePg�p-R"] Y?��ZXf9_PTMePZ p <3l� ]3YBT V&� K������v4 K�� �&b\#8m ��_PT s TQTos�_2sX_P[MsP_PT�ZXV�VrSPg s Rq[Ms"TaU

^\VrTQ[6R>V l�g _P]!� [!s-R-[QePg j T6RFg8T�YHT6s�[ Vr]!�3_ �Qs�_dZ g�YH]=R��=_=]=VX]9f3l�ZXg�U&ZXg s ZreP^6sP[6eOU2YG]3R"T��=R>ZkYGZrs"Z jPT!s�_P[MeHZre2s�_PT

V�Z jdZre&f;�4?'(��� ;�C 0ZreH�_PZX^o_$s�_PT�YHT6s�[\Vr]!�3_5Z�g�Y)SP^6_)YH]\R"T])<PV�Z �3S=T\4\Zks g�[�PZ�gD�=[\g8giZke=fs�_=RF]\S&f\_2s�_PT

YGTQU=Z p ]=gMg T6srs�T9m6� ]3R>T�^a]3YH�PVXTos T � ] R�s _)�pqR"Z�^6[Qe)Y�[6s"T6R"Zr[\V=]3p ^9]3Y>�=[QR-['<PVXT [afPTQ4QpqR"]=Y ���Xe2�$[QeOTQ^6_

�q��R"[QY$<=]\S&R-fP4&K��=Nvc3#�[ae=U���SDVr[ U �$[6YGZ�U3[��"AT6R"[6[ UPg [MePU � T6RqeO]9SPgig ZX43ZreH�PR"TQg g # ^6[6e5^6]\e&p Z�U=T6e\s"V l;<DT

RFTap"T6RFR"T\U s ]=�4?� 8�����3���.�!/���%'�! Hm ��_PT � ] R�s _)�p-RFZ�^6[aeHg��OTQ^QZ Y5ToePg�RFTMjPT6[\V9s�_PTEgL�PTa^QZXT\g p T'� UPZXg s-Zre=fvSPZ gF_PZre=f

^QR-[QePZ [ V3p"T6[6s�SPR"T9gi4Q[��$TQ[�A=�P]=gFs ]3R�<PZks-[ V=^Q]\e=g s�RFZ�^os"Z ]3e)[Me=U=�$ZXU=T!e=S=^a_=[QVP^6R>T6gFs"4Q[Qe=Us-_=TEgMVrT6eOU3TQR-ePTQg g�]3p Zks"g

YGT s�[��=]vU&Zr[\VXg6ma��_PT�Y [Ms"TaRFZr[\V=p-R"]3Y �$[\U3[aR�[ ePU ��Z9AOZ9A3[�R>TQg TaYH<PVXTQg �$]3R-s�_)�p"R"Z�^6[Qe ��?' 8�����K�����G/��9%��� .ZXe

[\VXVOs�_PTQg8T�p"T6[6s�S&R"T\gi49[ae=U)Z�g R>T6p"T6R"R"T\U2[Q^ ^Q]3R"U=Zre=f=Vkls�] s�_PT�g [aYBTg+�OT9^aZ�Tag6m

Page 11: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

���

$ '�K �/&%- �����%' '�J�)( �������%�� E ��ZXTos-R"Z�^o_d4&K��3LvJP43p�R>]3Y ��[QT6s ]=VXZ 4Q_P[Qg�[EgMZrYBZ�V [aRDYG]&Vr[QRdYB]3RM�=_=]PVr]=f9l

� ��ZXT6s�RFZ�^o_P4DK��\L=J't p"ZXf&m\J �PK �\L&bPt pFZrfDm ��� #Fma��]=R"T YH[6s T R"ZX[QVPp-RF]3Y ��[ T6s�]&VXZ��[Qg�UPTag ^QR"Z�<=TQU=<3l�A�S �QR"Z eH� K����vN3#

[\g��� ������������� ������! BD�+�� �%��'E&m � ^QR-Sdg _OTQU)gKA3SPV�V=�-A�S��aRFZred4dK����\NDt �PV-m�� m L&# Z�g�g Z YGZXV [aRDs ] s-_O]=g T�p-R"]=Y+s _PT

��]C��T6RD�1�2[ g _d4\TQg��PTQ^ Z [ V�V l;< l�Z s�gD<=R>]3[QU$e3S=^a_=[\V=^6R>T6g s"m A�S��6R>ZkePW g�[QgMgiTQR�s"Zr]3eH� K����\N't �=J3J=#Ds�_P[osdZrs-g�^6Rq[MePZX[QV

jP[6SPVrsdZ�g��6YH[QR�APTaU=Vkl�^ ]3e=^6[6j T��ZXg�e=]9sd^Q]\ePg ZXg s TMe s��$Z s _2_PZXg�%7Z fvS&R T �Dm L&�$m6��Z�g�U=Tos�[\ZXV�TaU$U=TQg ^6R>Z��=s Z ]\ePg�]3p

s-_OT e=S3YGT6R"]\SPg s�TQTMs�_>�=]=Zre3sd]9S3sPsX_dT6ZXR'g ZXYBZ�Vr[QR"Zks Z�T g �$Z s�_s-_O]=g T�p-RF]3Y �$[ U [aRD[QePU��Z9A=Z�A3[\m\��_OT�g [aYHT�Z�g

s-R-SdT�]3p s _PT S��!�OTQRDYGZ�V�A s�TQT6sX_HZ�V�V SOg s-R-[6s TQU8<3l��Z�Tos R"Z�^6_�� K��9LDbPt �PV-m KoN&4\p"ZrfDm �3JP# �6s-_PT!�2�dLHU=ZXp-p"T6RFg�pqR"]3Y s _=[os

] p��4?'(��O ;�� < ls�_dT�gF_P]3R�s"T6R5�=R>] s ]=VX])�v_)[Qe=U$YH]3RFT!s�Rq[aePg jdT6R>g8T YGTos�[\Vr]!�v_P49[ g�]3e$s�_OT YH[6s TQR"Z [ V&p�R>]3Y

�$[\U3[aR m ��_PT6RFTap"]3RFTQ4 $ '�J / %- ��9� %' '����(4 ���!� %���E�Z�g�^a]3ePgMZrU=ToR>TQU2[EgFl=e=]\e lvY ]3p5�4 ��+�5� �-�I�� ������!

8�����K�����!/���%��C Hm

��_PT Ta[QR"V�Z�TQg sPSve=U=]9SG< s TQU2R"T��=R>TQg8T6e3s-[Ms"Z j T ]3pPs-_PT �$p-R>Z ^Q[ae2�7V�Z ]v^6TaePT��4 '�+���������� ��K���! :^QV [aU=TEZXg s�_PT

p"]3RqY p"R"] Y.�![ae=f=T3<=[Q[ae��$T6f s�_=[Ms&��]v]=Z �iT6R'� K��3N3J=# Z e=^ ]9R"R"T9^Ms"V l�^Q[QV�V T UH�4?+B �K�� '%��'EDm ��_PT p V [Ms�s�T6ePZre=f$] p&s _PT

TQ^os ]=VX])�v_D4\V�]\e=f2eP[aRqR"]!�7�=R"]9s ]��D[ae=U2YGT6s�[\Vr]!�3_>�$ZksX_2s�ToePUvToeP^Mls"])^QVX]=giT sX_dT4�=]3g s�p"]=gMg8T6sXs-T 4 s�_PT�UvTQT�

V�ZrePf9S=[ V3]!�PTMePZ e&f�]3p=s-_PT YGTQU=Zr[ae2j=[ V VXTol 49[ g �$TQV�V3[Qg�s-_OT�V�Tae=f9s�_=T6ePZre=f$]3p=s-_PTE^ R"]=^M_=T6sP[QeOU2^aRFZ�g s-[E]3p=��7LP4

�_DZ ^6_)p-]=R"TQg _=[ U3]!�;s�_OT�^QVX]3g8S=RFTE]3p&s�_PT�YHTQUvZ p-]Pg giTos�s T�Z e$s�_PT�YH]3U=TaRqe)p ] R-Y � ��]P]=Z �iTQR"4 K��3N3J't �PV-maJ��3#F4Q[\R"T

['�=]3Y5] R��v_DZrT g�]3p&s�_dT �7VXZX]3^ ToedT#�� ������������� ��3���� .^\Vr[ U3T\4\Z e=U=ZX^6[as-Zre=fs�_=[6s=s-_OT ��[ae=f3T�<=[6[Qe���TQf2p ] R-Y

<PTQVX]9e=fPg�s�] s�_PZ�g�^ V [ U=T9m ��_=T�VX[Q^'A�]3pD^QR"Z�g s-[ 4)�$Ta['A3ePTQgig�] pd^6R>]=^o_PTMs"4Q[Qe=U�g s-R"[ Z f3_3s��=R"]3s�]PVr]!�3_�]9e$s-_=T

�&R"TaYH]&Vr[QR"g�[6R>T��=R"ZXYBZrs ZkjOT p"T6[as�SPR"T\g sk_&[Ms��)]\SPVXU �PTQR�_=[��=g �[QR"Rq[ae=sP[�g��OTQ^\Zrp"Z ^�e=[aYHT�UPZ g s Zre=^6sPp-R"]=Y

�4?) 8�����3�N���!/���%'�! Hm

�"eH]9s-_OT RP�[Qg sP�p"R"Z�^6[ eHg Zrs-T g 4!�4?' 8������� ���!/���%��� 0Z�g�]3p�s"TaeH^6[\VXVXT6U>��?-% '�K ��/6�)� �)�3� %��!/6�C �4'<3S=s=s�_PZ�g

e&[aYHTE� ��ZXV `a_dTQZrYBTaRF4&K��3JPb\#'Z�g <P[Qg8TQU2SG�P]9e)[E�-V�]=g s-#'gKA3S=V�V p-R>]3Y �V�U\S3jP[QZ!�_P]vg8T �wJH�-�Rq[aY$<P]3S=R-fP4

K��9L��Dt p"Zrf'maJ=L&#D_=[ g [ae)]!<PVXZ��9SPT�YGTos-[ VX])�=_P4\gMZrYBZ�V [aR&s"]sX_&[Ms ]3p s _PT�V�Z jdZ ePf$p-]=R-Y mQ�-s��PR"])<P[3<DV l�^\]3YHTQg�p-RF]3Y

s-_OT S��!�OTQR'V�TojOT\VXg�] pd�V�U3S3jP[ Z=����T6['AvT6l=4&K ���=bDt J�b\#>4'�_PTQR"T�U=T6p"ZredZ s T1��?'(��� 8�� .p"]=gMgiZ V�g [aR>T�A3eO]C�Ee

� ��]=]=Z �iT6RF4&K������3#8mQ��_=S=gM4��4?-%- '�� �G/��!� ������%3�!/��! :ZXg �=R"]!<=[�<PV l�g lveD]\e3l=YG]9SOgD�$Zrs-_$�4?'(��� 8�� �4 [ g!ZrePU3TaTQU

Page 12: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�M�

�Rq[aY$<D]\S=Rqf�� K �9L��dt �=c3c3#D_P[QU�g S=f3fOT6g s TQU)p-Z R"g s"ma�[Qg sd�pqR"Z�^6[Qe$�4?' 8������� ���!/���%��� 0Z�g A3e=]!�EeH<PTag s�<3ls-_=T

YH[6s T R"ZX[QV�4\Z eP^\V SPU=Z ePf)g8T6jOT6R"[QVPg�A=SOV V�gM4Qp-RF]3Y T6[ g sd[6eOU=�$T\g sd]3p=��['A=TE��S&R�A=[ae=[\46,�Toe=lv[\49YG]=g s Vkl�]3pdV [as-T

�7V�Z�]3^aT6e=T�[Qf=TE� �$[aRFR"Z�g 4 K��vN��=4&K�� � � �Q[aedU �OT6RFgMm\]�<Pg6m #Fmv��_=T6R>T�ZXg�[QV�g8]2[Eg�A3SdV V=p-R>]3Y s�_PTEVr[6s T � VXZ ]v^QTMe=T�]=p

E['�$ZX4 ,�T6e3lO[Q4vZke$s-_=T4��� ��Hm ��_PZ�g�YH[os T6R>Zr[\V=U=]=T6g�eP] sdUPZrpqp TQR�p"R"] Y.s-_=[Ms']3p=s-_PT!�$[\U3[QR&%7]3R"YH[os�Z�]\eP4

T�=^ T��9s=s _=[6s 49[ g eP]9s TaU=<3l�A�S��6R>Zke5� K����\N3#>4 s _PT YHTaUPZkp"]=gMg8T6sXs T�]3p&� ��Z g�S=gFS&[ V�V l�^\VX]=g8TQU=4\]!<3jPZX]9Sdg Vrl$[

U=T6R>Z j TQU)^M_&[aR"[Q^os T6R7VXZ e�AOT6U;�$ZksX_HZrs"g�lO]3S=e=f3TQRDfOTQ]=VX]3f=ZX^ [afPT9ma��ZOAPZrSPg���JPc3c\J&#D_=[ g eO] s"T U�s�_=Tg8[6Y5T

T6j=]=VrS3s�Zr]3e=[QR�l;�=[6sXs"TaRqeHZre2s�_PT!�7V�Zr]=^aTMeDT�]3pd� _P[ U&4Qp-R>]3Y ,�]=V�V�� s�],�] R"]$��] R ] ma��g�[6s=�$[ U [aR'[QePU ��Z9AOZ9Av[Q4

s-_OT e=S=^M_P[QV=^QR-T\gFsdZ�g@<PR"]3[QUvT RPs-_=[MeHZXe2s�_=T�V�Z jdZ e=f)p"]3R"Y � %7ZXf&m J=#F4'<=S=sd]9jPT6R-[\V�V=g ZX`QT�U=]3T g eO]9s'TO^QTaTQU s�_P[os ] p

s-_OT�V [Ms�s�T6R-<PT6p ] R"T!s-_OT�TQ[6R>V l2��V T\ZXg s ]3^aT6ePT9m

��_PT s�Rq[aePg Zrs"Zr]3es ]H�4?)( �9 ;�! �=RF]�<=[�<PVrls-]=]!A;�PV [a^ T�gF_P]9R-s Vkl�[Qp�s TQR&s _PT �7V�Z ]�� � VXT\Zrg s ]3^aT6e=T#<=]3Sve=U=[6R-l

Z e2�[ g sP�p"R"Z�^a[3m6�C]=V [aRFg�pqR"]3Y �V�U9S3jP[ Z)�$TaU �"�7� ��]P]3Z �8TQR"4 K������Dt �PV�maJP43p-ZXfDmaL-�qb��'�PV-m �P49p"ZXf&m��\#']=p s�T6e)_=[6jOT

V�]\e=f=�=R"]3s�]PVr]!�3_OgM4 [ e=U2[ V�V9s�TQT6sX_)p-R"]=Y �V�U3S=j=[ Z�4Q[\g �$TQVXV3[\g�s-_O]=g T�]3p=s-_PT!�>V�T6R>Tosdg�AvSPV V9,�� � � � �9J��v�

� �_P]vg T�[6fPTEZ g��=R"])<=[�<PVkl�7VXT\ZXg s-]v^6Tae=T3t��ER"]��e)[6ePU%'TaZ9<OT9VX4DK�����KvtXK��3#D_&[MjOT�YGTos-[ VX])�3_Pg �_PZX^o_�[QR"T YG]3R>T

]!<PVXZ �\SPT s�_=[aeHZre2s�_=T ��V�ZX]=^6Tae=T �4?' 8�������N���!/���%'�! Hm �!R>]3gMg��"g8TQ^os�Zr]3ePg s�_=R>]9S=fv_2sX_PT ��JH]3pd]3ePT�]3p&s-_PT g T

TQ[6R>V l)R>T��=R>TQg TMe s�[6s Zrj=T g�]3p���?'(3�� ;�! :pqR"]=Y+�7TaePZke����"AT6R"[6[ U=gM4dK����\N3#'g8_=]��Cs _=[6s=s�_PZ�g�ZXg eO]9sP[�R"T\gFSPVksd]=p

s"]v]9s�_>�$T [aRit6[ Vks-_P]\S=f3_2sk_PT�V�]��3_Pg <=Ta^Q]3YHTEg s�R"[ Z f3_=s�TQR-�$Z s-_>�$T [aR>4 s�_dT6ZXR']9jdT6R"[QV�V=])<dVXZ �\SPToeOTQg g�Z�g�eP]9s

[\V s"T6R>TaU)� %�Z f&m��=#Fm\��_=T#�=R"]9s-]=VX]��=_HT3s TMePU3g <PTolO]3e=U2YGZXU!�-V�T6ePf9s�_)]3p=s-_PT!s"]v]OsX_d4Q[6ePUsX_PT [3PZ�g�]=p s�_dT

YGT s�[\V ]!�3_H�=[ g giT6g�s-_=R"]\S=f3_2s�_PT Y5T6U=ZXp"]=g g TMs�s T9m

�"e2��]3R�s�_)�p"R8ZX^Q[Q4��4?) 8�����3� �I��/���%'�! +Z�g@�=R>T6giT6e9sdp-RF]3Y s _PT�Vr[6s T � VXZX]3^QTMe=T�S=e3s"ZXV3sX_PT�Vr[6s T

�7V�T Z g s ]=^6TMeDT\m6��sd��_PV3[ V=��S&fv_PV [aY Z e��]=R-]=^ ^6]P4\U3[6s�TQU�s ])^Q[9m JDmrb���[E�qAT6R"[a[\U=g�T6sP[QV-mn4&K���� �\#>4 s _PT s�T6T6sX_d4

�_DZ ^6_)[aRFT�[QV�V=U=TQ^QZXU9SO]3S=gM4Q[6R>T�Z�U=Toe=s ZX^6[\V9s ] s�_P]3g8T]3p=�)[\U3[aR'��� � �������$]=R&,��0� � ��J��9J=c$pqR"]=Y+s _=T

I;���=T6R-�ES=R"f=Z9� T6YH<dT6RD[6s=,�]v]!<PZ9% ]3R-[3m\1 �OTQ^QZrYHT6ePg�p"RF]=Y ��SDVr[\U �$[aYGZXU3[�ZXe2� ]3R-]=^Q^ ])�-ATQR-[Q[QU=g�[ae=U

��T R�eP]9SPgig g ZX4vZke>�&R"TQgig #"4\U [Ms T6U s�] s�_=TEg TQ^a]3e=U_=[\Vrp']=p&s-_OT YGZXU3U=V�T!�7VXTQZXg s ]3^\TMePT�� �_P]vU3T\g�T sd[QV-m 4DK������3#D[ ePU

Page 13: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�M�

pqR"]=Y+s _PT�VX[os T ��V�TQZXg s ]v^QToedT����ERF[aY <d]9S=R"f=4&K���� �dt �dV�m)�=49p>Z fPm J �'�PV-m N #"43[QR>T�^ V�T6[QR"Vrl)R"TQp T R-[�<PV�T s ] s�_PZ�g�gK�=Ta^9Z T9g

[Qe=U$e=]3s=s-]sX_PT�YH]3UvTQR-e)p"]3RqY m

��_&[ae=f=Tag�Z eHg8Z ` T�[Me=U;�&R"]!�O]3R�s ZX]9ePg�]3p s _PT YGT6s�[��O]=U=Zk[ V g�R"T\p-V�TQ^6s=s-_=T�Toj ]=VrS3s"Zr]3eH]3pdV�]=^ ]3YH]9s ]3R

[\U3[3�=s�[as-ZX]\eDg�Z e2s�_PT#��?' 8������� �I��/���%��! �-�4?)(3�� ;�! :V�ZrePTM[6fPTE� % ZkfDmaL=#8m\��_P]=g8T]3p=s-_PT�TQ[6R"V�Z�TQg sPp"]3R-YGg�p-R>]3Y

��[Qe=fvT�<=[Q[Me��$TQf$]3R'pqR"]=Y+s _=T!,HTMe3l=[aeHg Zrs T6g�] p=� �PTagMZXU3[Q4v�!_=T6YHTQR ]3eP4 [ ePU2� s�T6RFZrR>49[QR"T�R"[Ms-_PTQR'g V�ToePUvT6R

[Qe=U�]3pDYHT\U=ZrS=Y g Zr`QT�� V [Ms"T6R&� VXZ�]v^\ToeOT p"]3R"YBgi43p-RF]3Y ��_PV3[ V=��S=f=_PV [aYB46��Z�A=Z9Av[\46�$[\U3[aR84Q[QePU ��[\Tos ]=VXZX49[aRFT

V [aRqfOTQR-<=S3sd]3p'gMZrYGZ�V [aR-�PR"]!�P]3R�s ZX]9ePgMm\1DZr`\TEU=ZXg s-R"Z�<3S=s Z ]\eHZXg eO]Osd^aV�Ta[QR"Vkl;<PZXYB]vU [ V=Zre2s�_PT�VX[os�s�TQR'g Zrs"TagM4

g S=fvfPT6g s Zre=f s�_=[6sdT Zks�_=T6RG" ��%' '�K�!(�[6ePU>� '���C����� �� ��3���� 0�)TaR"T]3pDgMZrYGZ�Vr[QR-�=R>]!�P]3R-s-ZX]\ePgM49]=R&s�_P[os�" ��%� ��K�!( Z�g

['<Pg8T6e3sdZre2s�_PTQZ RdYBT6s�[��=]3UPZr[\V=g [aYH�PVXT9m ��_PT R T�Z�g eO]2U=]\S!<=s-4\_=]!��T jOT6R>46s-_=[6s=s _PT6giT YGTos [3�P]3U=Z [ VXg YG]vg s"V l

RFT��=RFTag TMe s��4?' 8�������N���!/���%'�! Hm ��_PTET6[QR"Vkl��V�TQZ�g s ]3^\TMe=T!YGT s�[��=]=U=Zr[\V�g p�RF]3Y �VXU=S3j=[\Z3[QR"T�RF]�<3Sdg sP[ae=U

gMZrYBZ�V [aR'Z e>�=R"])�=]=R�s"Z�]9eOg s�] s�_=]3gMT�]3p=s-_PT�YB]=UvT6R"e)p"]3R-Y mQ�*ZXU=U=VrT�[ae=U)VX[os T ��V�TQZXg s ]v^QToePTEg+�PTQ^QZ Y5ToePg�p-RF]3Y

�[\g sd[ae=U � ] R�s _)�p-RFZ�^6[E� s�_dT 1P]\S3sX_)�p"R"Z�^6[ e p ] R-Y YGZ�fv_3sPRFT3�&R"T\g8T6e3s�"$��%� )�K�!( #D[QR"T�gMVXZXf3_=s-Vrl�g YH[\VXV�TaR m

��_=S=gM4\ZksdZXg!VXZ�A=T\V ls-_=[6s=s _=T Y5]3UPT6R-e)p-]3R"Y U=T6R>Zkj T6U2p"R"]=Y+s-_OT��V�U9S=j=[ Z&]\e=T4< l�giZ ` T R>TQU3S=^Ms"ZX]9eP4��_PZ�VrT!s�_OT

��] R�s _)�p-R>Z�^6[ae=�P]��3SdVr[6s Z ]\eDg�R"T6s�[\Z ePTaU s�_PT�g VXT6eOU3TQRDYGTos�[��P]3U=Z [ V�g�]=p���?� 8�-�G��� � �G/���%��� Hm ��_PT6RFTap"]3R"T94'<O]OsX_

s-_OT�U=TMe3s"Z s Z ]\ePg�[Me=U;�P]3g s�^QR-[QePZr[\V�g�gFS=f=fvTQg s=s _=[6s=s�_=T!s-R-[QeOg Zks�Zr]3e)p-R>]3Y �4? 8���!�J�����!/���%��C *s"];�4?'(��9 ;�! s�]=]!A

�DVr[ ^6TEZ e2�[ g sP�p"R"Z�^a[3m

AT6e3SPg-" � %) ��K�C( A�R"[olO4�K �QJ�K

7('*) � �+) �-, � �1�J�O"$��%' ����!( F!��%�� �+/���(���Z e=e=[ T6SOg 4 KoN&b��v4vV ZrjPZXe=f2�p"R"Z�^6[ e/.�<DVr[ ^�A#03 E_PZre=]&ma��]!�$TQR

�7V�T Z g s ]=^6TMeDT!s�] �Ta^6T6e3s�4Q�pqR>Z�^6[vm

��� �12�������� �3� �7R"T YH[3PZ�V�Vr[�[�<Pg8T6e3s ]3R&jPT g s"ZXf=Zr[\V-mQ�!R"[6ePZrS=Y?g _P]3R-sP[6ePU2R"T\Vr[6s Zkj TQV l;<&R"]3[ U&m

��T6S=R>]3^6R"[6edZ SPY+s�Z Vrs T6U2[ e3s�TQR-]=U3]3R>g [ V�V l)R>TQVX[os�Z jdT!s�] s _=T�gK�PV [MeP^M_3eP]v^6Rq[aedZ S&YG4QR>TQgFSDV s"Zre=f�Zre)[�YB]3R>T

jdT6R-s-Z�^6[\VXVkl�]3R>ZXT6e3s T6U�]3^Q^QZ��PZrs�[ V)�dVk[ae=TE]3R7TojOT6eH]9eOT�ZXe=^\VrZre=TQU$[6e=s�TQR"]3U=]3R>g [\VXVrl=4\[�e=SP^6_=[QV=^aRFTQgFsdV�Tag g

T�!�=[ae=U=T U �O]3g s"TaR"Z ]3R>VklO4\[ YG]=R"T�UPT6T3�dV l�^ ]9ed^6[oj T�^6R"[ae=Zr[\V!�=R>]3p"Z V�TQ49[1<=[QgMZX]v^6^\Z��dZ s-[ V [ae=fPVrTQU2R"T\Vr[6s ZkjOT!s"] s�_=T

Page 14: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

<&[ g ZXgK�3_=Tae=]=ZXU=49[Eg _O] R�s�ToePTaU2p-[ ^\T��$Z s-_H]3R�<PZrs g YH]=R"T [ae9s�T6R>Z ]=R"Vkl;�P]=giZks ZX]9ePTaU2[aePU2^QVX]=giT6R&s"]s�_OT e=[\g8[\V

eP]9s ^o_P4\[6eOU)] pXs"TMe)[�e&[ g ]=Vr[\^aRFZrYH[\V=^Q]\e3s-[a^6s"m

"$�N%� ��K�)(�B �+�� '%���E�� �H]v]=Z �8TQR�� � [os�s"T6R>g ]\eD4&K��=N\J=#

�4 '�+�5�������6 '�����! B ���� '%��'E�$]=]=Z � T Rd[6edU �'[Ms�s T6R>g8]9ed4dK��=N9J't K��

7('*) �'���7]=]3R"Vkl;�=R>TQgiT6R�j TQU)Z ed^a]3YH�PVXT s T�g�A\SPVXV p-R>]3Y+,�[6e=[��=]=Z�46,�T6e3lv[\4Q,��0��,� ��� �)� ��]&]=Z �iTQRD[aePU

�'[6s�s-TQR"gi]9eP4 K��3N3J't p-ZXfDm �3�#8m

��� �12�������� �3����_=T�]3R>ZkfPZreP[QVvU&Zr[Qfve=]=giZ g�^a]3e=gMZXg s�g!TMe3s"ZrR"TQVrlH]3p-�PV�TQg ZX]\YH] R��=_PZX^E^o_=[QR-[ ^os T6RFgMm\��_PZXg

g��PTa^\ZrTQg�_=[\g�]9edV l$[�p"T��.[��D]3YH]3R��v_DZ ^ p"T6[6s�SPR"T\gD�$Z s�_)R>TQg+�DTa^6s=s�])Zks�g�VXZ�A3T\V l�[6eP^QT g s�]3R ��?'/� ��! ��2 �3�"t ]3R�<DZ s

YG]3RFT�[6e3s T6R"Z�]3R �$Z s _)RFT gL�PTa^6s=s�] s _=T s�]v]OsX_�RF]!�$4vg�A=SPVXV!�3R>]3p"ZXV�T YG]3R>T�^a]3e=^6[ajPTQ4v]3^Q^QZ��PZrs�[ V)�PV [MedT Y5]3R"T

jdT6R-s-Z�^6[\VX49[ae=U$e=S=^M_&[ V3^QR"T\g s VrT\g8g�T�3s TMe=U=TaU=�=]=gFs T6RFZX]=R"VklPm

�� ��2���� �[QR"Vrls"])Vr[6s T � VXZ�]v^QTMePT946�[\gFsP�2p-R>ZX^Q[9m

��� � ,)� � ) �� ���9���� �8, ����� � ��� ��� "5����)�:� ���N�q1�A=SPV�V3� � � KaJ�����J�KE�q13�@Ka#'Z�g <=RF]!AvT6e5Z e s ]

gMTojOTQR-[\V!�PZXT6^\Tag6m6�0Vk[ R-fPT�p-Rq[af=YGT6e3s'^a]9edg8Z�gFs-g�YH[QZXePVkl�]3pDs�_DT�]v^6^\Z��dZ s-[ V [ae=U8�&[aR-sd]3p s�_dT�^6Rq[aePZ [ V�<=[agMT3tQZ s Z g

s"]v]2U3[QYH[afPT6Us-];<=T p-SPV�V l�U&TQg8^QR"Z9<OTQU&ma��_PT�]9edV l�^\VXTQ[6R'p>T6[6sXSPR"T�Z�g s�_PT�g _=[aRM�Hg [QfOZksXs-[ V�A=TQTQV�4Q[os VXT [ g s�Kac$^QY

V�]\e=fP49]3es�_=T <=[ g Zr]P^a^9Z �PZks-[ V�mo��_DZrg APTaT9V=Z�g [ V�g8]2p"]3SvePU)Zre$�4?)/6 '�! � 2 �J�=[6eOU�"$?�F!��%'�G��/���(a4'<=S3sdZksdZ�g Y)SO^6_

�$TQ[�AOT R Z e$�4?' 8�����J�����!/���%��C [Qe=U>��?'(��� 8�! Hm � _OT!e3SP^M_=[\V=^6R>T6g sdZ�g�U3T6jPTQVX]��PT6U2Y5]3RFT4�=]Pg s T6R>]3VX[os�T6R"[QV�V l

s-_=[aeHZ e "$?6F!�&%�����/6�9(a4�<3S3s=s�_=T]v^6^\Z��dZ s-[ V&ZXg@�PR"]!<=[3<DV l�V�TQgMg�ZreO^QV�Zre=TaU=�P]vg s TQR"Z�]3R>V ls _=[QeHZ e

�4?) 8�����3�N���!/���%'�! Hmo� _PT4<=[\g8T�] pPs-_PT�]3^Q^ ZO�PZrs�[\V!�[Qg�e=]9sPYHSP^o_;<PR"]3[QUvT RPs-_=[Me2s _=T s ]��D4\Z e5^6]\e s�R"[Qg s=s ]

"$?�F!�N%����K/��O( m6�"eHg SG�=TQR"ZX]3R&jdZXT'�$4 s _PT�g [af=Zrs s�[\V3eO]Os ^6_H<=TMs&�$TaT6eH<O]Osk_H�$Zre=fPg�]3p s _PT e=S=^M_P[QV=^QR"TQgFs'ZXg

g _=[ V�VX]!�)m3�$TQVX]!�;sk_DT�^6R>TQgqs"46s-_PT�YHTQU=ZX[QV��=[6R-sd]3p&s�_dT�]3^a^\Z��dZ s [QVPZXg�UvTQT��PVrl$_=]=V�VX]!��T6UP4Q[Qg!Zre$�4?�(3�9 �! Hm\��e

s-_OT��_P]=VXTQ46s-_OT ]=^ ^QZ��=Zks [QV=Z�g�gMZrYGZ�VX[6RPs"] sX_P[Msd] p��4? /� '�� =� 2-�3�=]3p&s�_dT�Vr[6s T �CZX]3^QTMe=T3m

��ed]9s�_OT6R5�PZXTQ^6T�^Q]\ePgMZXg s"g�]3pPYG]&gFs']3p=s-_PT�YH[dZXV�VX[E� %�ZrfDmvb\# m ��_PT�<=[ ^'A$]3p s _PT eP[Qg [ V9eP]9s ^M_HZ�g

['<=]9j T s�_PT R"TQ[aR']=p&�'JP4 s�_P[osd]=p=s-_PT�ZXe=p-R-[\]3R�<dZ s [QVPp-]=R-[QYGT6e)[�<P]9j=T!s�_=T R>T6[QR ] p&� �=49[aePU s-_OT p"R"] e\sd] p=s-_PT

Page 15: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

���

] R�<PZ sPR>T6[\^M_=T\g�s�_OT�V�TojOT9Vv]3pD� K\m ��_PT6giT��O]3gMZ s"ZX]9ePg�[QR"T�YG]3R"T [ae3s T6R>Z ]=R&s _=[MeHZ e)[ V�V=gK�=TQ^ Z T g�]3p

�4 '�+�5�������6 '�����! .[aePU)^QVXT6[QR"Vrl�g _P]!�;s�_P[os=s�_PZ�g�gK�=Ta^9Z T9g�_v[\U2[!s ]=]9s�_)R"]C�*YG]3RFT4�O]&gqs"TaRFZX]3R"Vkl�gMTMs"4Q[Qg�Z e

"$?�F!�N%����K/��O(o49[Ms�s T\gqs"Z ePf s�_P[os=s�_OT�g�A\SDVXV3R>T6[QR-R"[6ePf=TaYHT6e3sd]=p s _PZ�g�V ZXe=TQ[af3T�_=[aU$[\VrRFT6[ U9ls-[�A=TQeH�PVX[a^QT�� %�Z f&m �3# m

�E]Osk_2s"]v]9sX_)R>]!�$g�]=pD� � � KaJ�����J�K�[aRFT#�$TQV�V��=RFT g T6R-jPTaU [ e=U2YG]3U=TaRq[Ms"TQVrl;�$]3RqeP4\T�O^QT3�3s&� �P4

�_DZ ^6_HZXg�T g g TMe3s"ZX[QV�V l$S=e��]3R"eP49[ae=U ��!K 4'�_dZX^6_HZrg!g8]=�$]3R"e2s�_=[MsDeO]9ePT�]3p'Zks�gD�=[Ms�s T6R"e)R"TQYH[QZrePg�jPZ�gMZ�<=V�TOm

��_PT!_=T\Zrf3_3sd]3p&� �)ZXg �DK�YHYG4Q[6ePUsX_DT!s�TQTos�_)[aR>T s�_PT6R>Top"]3R>T��=R>]!<=[�<PVrl�g V�Zkfv_3s"V l$YG] R"T�_3l��Og ]3Uv]3e3s=s-_=[ae

s-_O]=g T�]3p!" ?!F!�N%����+/���(\m6��V�V3s"TQTMs�_�[6R>T�giZ YGZ�Vr[QR&s�] sX_P]=g8T�] p��4?)/� '�� � 2 �3��4��$Z s _)[Eg s�RF]\e=f)^QR"]=^M_=T6s"4'<3S=s=eO]

^QR"Z�g s [9m ��_=T4�PR"]9s-]=VX]��=_HZXg eOT6[QR"Vkl$s�Rq[aedg jdT6R>g8TQ43]\ePVkl�gMVXZ fv_3s"Vkl�^MS=R-jPTQU)]3R T9g8giToe3s�Z [ V�V l�g s�Rq[ Z fv_ sd]\e2s�_=T

�&R"TaYH]&Vr[QR"gM4\UPZrg s Zre=^6s VklH1��"g _=['�OTQU$]9e2s _=T Y5]=V [aRFgi4)<3S3sPs�_DZ ^ A$R"T\Vr[6s ZkjOT s�])Zks g�V TMePf9s�_Dm � _OT��=R>]9s ]3^Q]\e=TEZ�g�g s-Z�VrV

RFT ^Q]3f\eDZ `\[3<PVXT [ g�[EU=Z�gFs"Z eP^MsdT6e9s�Z s�l�]3e2� �2[ae=U�'LDm � _OT YGTos [QVX]��=_HZXg�gMVXZ f\_3s Vkl�]!<dVXZ��9SPTQ4'<=S3sdg _O]3R�s�4'<=T\Z ePf

V�Z s�s�VXT YH]=R"T s�_=[ae)[�p V [Ms�s"Tae=TQU)^MSPg���]9e$s�_PT�YH]PVk[ R"gM4 [aePU$Z s�g�V�Zre&f\S=[\V!�=[QR�s ZXg�e=]9sd^MS=R-jPTQU=�O]3g s�TQR"Z�]3RFV lDm �

^Q]3YH�PVXTos T�^ Zke=fvSPVrS=Y g S=R"R>]9S3ePUvg s _=TEV�Z ePf\S=[\V=g ZXU3T]3p=s-_PT#�=RFTMYH]PVr[QR"g �\ZrsdZ�g �$T6['A=TaR�]3e$�DL'mM��]!�$T jOT6R>46s-_PZ�g

^\Z e&f\SPVrS=Y ZXg!Vr]��$4Q[6ePU s�_PT4�=]=g s�p"]=gig T6sXs TQg�]3p&s-_PT�YH]=V [aRFg �$]3SPVXU eO]9s'^ V ]vg T4<PT6p"]3R>T�Vr[6s T��$TQ[aRF4 �iSPg s�<=TQp-]=R"T

s-_OT9ZrR'UPZrg [����=T6[ R-[Qe=^aT3mQ��_PTEVr['<=ZX[QVPg ZXU=T�]3p&�'JHZXg!TMjPT6ePVrl�^Q]9e=jOT� 4'<=S3s=s�_O]=g T�]3p&� ������J�[6R"T�YG]3R"T�gMZrevS=]3S=gi4

�$Zks _2[�U=Z gFs"Z eP^Ms��&[aRq[ ^a]3e=T p"]=V�U)Zre2s�_=T�S��!�=TQR-�=[ R�s']3p=s-_PTE^ R"]!�eP4'�_dZr^6_H��]9SPVXU jO[QePZXg _5Z eHVr[6s�TQR-�$Ta[QRFm

��_PT4�=[QR-[\^a]9edT p-]PVXU)ZXg�gFs-R>]9e=fOT6R']3e2� �P4'�_PZX^o_)_P[ g [!s-R>Zr[Qe=f3SdVr[QR']\S s V�ZnePT9m ��_PT ]v^Q^QVrSPg [ V=g S=R"p�[\^QTE^Q]9ePgMZ g s-g

] p=sN�$])UPZ g s Zre=^6s6�PV [aePT6gM4\Z eOU=ZX^6[6s Zre=f s �])U=ZXg s Zre=^6sd^o_PT��Z e=f8�v_&[ g T g 49[aePU sX_OTVr['<PZ [ V!�[\VXV�g�]3pDs�_DT

�PT6R"Y)[Qe=Tae3s=s"TaT sk_2_P[ojOTEg _=[QR��H^aS=g+�Dg�ZreHVr['<PZ [ V9jdZrT'�)m

� ��1� * )���,)� � �1��� � � � � & � �������-��e�S����=T6RDs ]=]9sX_)R>])�$43��� � ��� L-� KE�F13�$L=#>4\Z�g�[QVXYB]=g sdZXUvToe=s Z ^a[\V

Z eHgiZr`QT�[Qe=U2Y5]3R �v_P]=V ]3f\ls"]2��� � KaJ � �XJ�K 4\T� ^6T��3s=s�_=[as=s-_=TaRFTEZ�g�[�g8YH[\VXV&^6R>ZXg s�[�]\e2��JDmM�*]3g sd]=p=s�_dT�]9s�_PT R

g��PTa^\ZkY5T6e=g�^Q]9eOgMZXg sP]=pDZ�g ]=V [Ms"TaU s TQTosk_H] Rdf3R>]3S��=g�]3p&sN�)]$] RPs�_PR"T\T s�T Tos�_2s-_=[MsdU3]2eO]Os U=Z p-p"T6R7YH[6RMA=TQUPV l

pqR"]=Y+s _PT�]3e=TQg�U=T6gi^6R>ZO<PTaU2['<O]9jOTOmo��_DT�[Qgig ]=^ Z [Ms-TaU ��J$[6edU � �P4Q� � � ��K���� K\4Q[aRFT�[QYH]3e=fs-_=T

fPT6]=VX]3fPZr^Q[ V�V l$lP]3S3ePfvT6g sd]\ePT6g��"g S�<��-YBTMYH<PTQR&��$J=#8m ��_PTQZ R'VX[6R"f=TEg ZX`\T�� � J't1�3J=5N9L)YHY �Q� ��t N=c 4���

Page 16: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

YHY # Z�g�]3S s�gMZXUvT s-_OT R"[6e=fOT�]3pdgMTojPTQR-[\V9s�ToePg�]3pdg��PT6^\ZrYHT6ePg�]=p)" ?GFC��%'���+/���(E�-��[�<PVXT ���\A�S��6R>ZreD4&K���� cDt s-[�<PV�T

b3#Fm

� � S3j T6ePZXV TEg�A\SPV V=p-R"[6fvYBTQe3s�4 ���#� KaJ���� JPc��q13� KQ#"4v^Q]9edg8Z�gFs"g�]3p-<P]9s�_2s�]3] s�_)R"]��$g �$Zks�_2��!K�� �$�DL

Z e)Y5T6UPZ S=Y*�$TQ[6R �qR"Z fv_ s=��!K YBZ�g8gMZ e&fv#D[QeOU=<=]3sX_$S=e��]3R"e2�.Kag�� %7ZXf&m N=#"49s�_dT R"ZXf3_3sP`6lvfP]3YH[6s ZX^ [aRF^a_D4Q[aePU

s-_OT fPVXT6e=]=ZXU2p ]&gig [9m ��_PZ�g�g+�PTa^QZ YHTMeH^Q[Me;<PT R"TQp-TQR"RFTQU�^Q]9e=p"ZXUPToe3s Vkls ]�"$�N%' '�K��(�<3ls-_PT�^6_=[aRq[ ^6s-TQR"g�]=p �:K\m

��_PT4�=[QR-[\^a]9edT p-]PVXU)ZXgD�$Ta['A=TQR&s�_P[MeHZre "$?GF!�&%'�G�L/���( 4Q[6ePU sk_DT6R"TEZXg�[Eg V�Zrf=_3sPYHT6s�[ ^6]3ePT�p"]=V U$]3e2��dJDm\�VXV

YGZ�V�A s�T6T6s�_)_=[MjPT�[�VX]3e=f�^6R>]3^6_OTosD[aePU[^6R>ZXg s�[ 4 <vS s=s�_OT [ae s�TQR"]��&�P]3gFs-TaRFZ�]3R"Vkl�T\VX]9ePf3[6s�T U2YGT6UvZ p"]=g giTos�s T

�$]3SPVXU eO]9s'^QVX]vg T�ZreHVX[os�T6R-�$TQ[aRF4\T�P^aT��3sd]3e2��dJDm\��_PZXg�s"]v]9s�_)_=[Qg�[EU=]9S�<OV�T�^6R>Z g s [Q4Q[QePU$[ YGTos [QVX]��=_

TQYHT6R"f=Z e=f8�O]=g s T6RFZ ] R"Vrl)p-R>]3Y s�_DTET6^6s�]PVr]!�3_P4vg8] s-_=[Ms&sk_PToR>T�ZXg�eO]=�=]=gFs-p-]Pg g8T6sXs TOma��g�]3e2s�_OT��PT6R"YH[6eOT6e3s

s"TaT6sX_D46s _=T YGTos [QVX]��=_HZXg�gFs-R-[\Zrf=_3s"m\��_OT R"TEZXg eO])Zre3s-TaRqe=[\V=^ Zke=fvSPVrS=Y m

� ����JPN3L���JHZXg�[�p-R"[6fvYGT e3sd]=pPY [3dZXV�Vr[8�$Z s-_2�� J�� �� �)�"g s-R"[Ms"Zrf=R-['�=_PZ�^EVrT6jOT9V3Sve A\eO]C�Ee=# m ��_=T\g8T

s"TaT6sX_)[QR"TEgMZkYGZ�V [aR&s"] s _=]vg T�]3pd� ��� KaJ ����JOc)ZreHVr[\^�A=Zre=f$[#�D]vg s�p"]=gMg8Tos�s T ]\e2��'J'ma��_dZrg YG]=R��=_=]vV ]3f3l�ZXg

V�Z s�s�VXT ^a_=[Qe=fvTQU)Z e)Y5]=U=T6R-e "$?GFC��%'�G�+/-��( 4\T�=^aT��3s=s _=[6s=s�_=T6R>T�Z�g�]3p�s-TQe)[aeHZre=^\Z��PZXT6e3s��P]vg s p"]=g g TMs�s�TE]9e2��'JDm

I;���=T6R'YGZXV9As T9TMs�_�]3p�" ?LB ���� �%'�'E�^ [Me><=T�U=ZXg s Zre=f3SPZXgF_=TQU$p-RF]3Y s _=]=g T�]3p��4?! 8���������.��/��9%��! <3ls _=T

g s�RF]\e&fvT6R5�=[aRq[ ^\]9e=T�p"]=V�U=4vg s-R-[ Zkf=_3sPY5T s�[\V ]!�3_)eO]Os ^6]3e3s�Z e=SPZre=f)Z e=s�] s _=T�UPZ gFs-[ V=^\Z e&f\SPVrS=YG43VX]3e=f$[MePU)])�OT6e

YGTQU=Z p ]=gMg T6srs�TQ49[aePU2p-R>T �9SPTMe3s Vk[ ^�A$] p��P]=g s-p ]=g g8T6sXs-T 4)<3S3s=s�_dT�U=Z gFs"Zre=^Ms"ZX]9eHZXg�e=]9s T6[ g l;�$Zks�_P]9S3sdT�=[ YBZnePZ e&f

s-_OT [\^os�S=[ V&g+�PTa^QZ Y5ToedgomM�-s Z�g�s-_PT6R"T6p"]3R>T�U=ZXp-p"Z�^6SPVrs=s"])ZXUvToe=s Z p�l$s�_PT sN��])��YG] ��'JPg�Z�VXVrS=g s�Rq[Ms"TaU=<=l��]3]=Z � TaR

� K��vN&b\#F4'�_PZX^M_HU&Zrp"p"T6RDYG[QZXePVkl�Z e$s�_PTEVr[\^�A�]3pD[E^QZ e=fvSPV S&Y?Zre2s-_OT�g��OT6^\ZrYHT6e�R"T p TQRqR>TQU8< l$_PZrY s"]

�4 '�+�5�������6 '�����! Hm6��]!�$TMjPT6R>46s-_PT [�<Pg8T6e=^ T�]3pP[^\Zre&f\SPVrS3Y ]\e)� � � KaJ ����JOc)gF_P]!�g�s-_=[6s=sk_DZrg�Z g�eP]9sP[

RFT V�Zr['<PVXT p T [Ms-S=R>TQ4\[6eOU s-_OT���YH] ��7JOg [aRFT#<=T6sXs T R ^Q]9ePgiZ UvTQR"TQU2[ g ��Z�^6TQR"]3s�ZrePZ=Z e=U=TMsFm

��_PT YH[MePU=Z�<PV T�� � � K J3L��"N9JH�&����#D_=[ g�[�^ ]3e=U9lPVXT#��_dZr^6_HZXg!Z ed^aV�Z edT6U$YHT6UPZr]�� jPT6e3s R-[\VXVklO43[ g�ZXe

"$?�F!�N%����K/��O(o43ZreH^Q]\e9s R-[\g sPs ] s�_=]Pg T�]3p �4?'(��O ;�� [Qe=UH��? ;�������N���!/���%'�! +����� � KaJDbG� Ka#Fm6��]!�$T6jOT6R>4 s _=T

YH[Qe=U=Z�<PVXT6g�U=]e=]9s U=Z g+�PVX[ol�[6e3l�]3s�_DTaR'g��OT6^\Zrp"Z�^�p T [Ms-S=R>TQm

Page 17: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

"$�N%� ��K�!(#F!��%��!�K/��9( � ��Z e=e=[ T6SOgM4dKoN&b��\#

��� �12�������� �3�q1dZ `\T�g8YH[\VXV TaR&s-_=[Qe1"$?+B4�+�� '%���E&mM%7[Q^aT YG]=R"T [Qe=fOVXT6U$]3e2s�_PT!ePT6S=R-]P^6R"[MePZkS=YG4\e3S=^a_=[\V

^QR"TQg sPeP]9s T�G�=[QeOU3TQU&m ��_=T T As�TQTMs�_�e=[QR-RF]!��TQR"4 [ ePU)V�]��3_Pg�YB]3R>T s�R"[6ePgFjdT6R>g T\m6�'R>T6YH]=VX[6R'R"]!�.g _O]3R�s T6eOTQU&m

�� ��2���� �[QR"Vrl�7V�TQZ�g s�]3^QTMePT s ]� T ^6T6e3s 4Q�[Qg sP[QeOU 1O]3S3s�_��p-RFZX^Q[9m

�� ���)� �:� � "3���� � � ���N� �"e2s _PT � ]!�$T6RD�8�[\g8_P49YG]3g sd]3p=s"_=T�Rq_PZre=]$Y [as-TQR"ZX[QV&p�RF]3Y s�_dT�T6[QR"Vkl�YGZXU=U=V T

�7V�T Z g s ]=^6TMeDT�]3pD��g�<P]&V T��"AT6Rq[a[\U=g�TosP[ V�mn4vZke>�&R"TQgig #DYGZ fv_3sPRFT3�&R"T\g8T6e3s=s�_PZXg�gK�=TQ^\ZrT g 4)<vS3sdZksdZ�g s�]3]

Z eO^Q]3YH�PVXTos T p"]3R'U=TQp Z ePZks�T�g+�=T ^QZ p"ZX^�Z�U=T6e9s�Zrp"Z ^Q[Ms"ZX]3eDm

� ���/)����:� �-�����

�V�V9�[ g s'�p-R>ZX^ [ae)R"_PZ eO]3g�R"T6p"TaRFRFT U s ]�"$��%' ��J�!(�_=[\U8�=RFT jPZ�]9SPg Vrl;<PTQTMeHZreP^QVrS=U=TaU)Zre "$?GF!�&%����L/���( 4

U=TQg+�DZ s"T s�_OT p"[Q^6s=s�_&[Msd]=^a^6S=RqR"TMed^6T�]=pD[6e5T�vs-[Me3s g+�=T\^6Z�T6g�Z e$s�_OT�TQ[aRFVXZ�Tag s=�7V�Zr]&^aT6eOT����;A=]9ePUv] % ] R-YH[Ms"Z�]\e)]3p

I$f=[6eOU [ 4\A�S �aRFZ ed4&K����\L=# �$]3SOVXU=�=TQR-_=[��=g <PT S3eOZ��9SdT [MYG]3e=f2YH[aYHY [ V g6mo��_PT�g�A\SdVXV ��� � KaJ ����J'KEZXg

gMZrf3ePZ p Z�^6[ e3s�<PT6^6[aSPg TEZ s'VXZre�APg s ] fvTos�_=TQR&s�]3]9s�_)YG]3R��3_O]&V ] f\ls-_=[Ms-�[ g R>TQp"T6R"R"T9U;�=RFToj ZX]9SPg Vrls ]�"$?�F!�N%����+/��9(

�$Zks _2[Qe�]3^ ^QZ��3S3sPs�_P[MsP^Q[6eveP] s�<PTEZrU&TMe=s�Z p Z�TQU2[ g�gFSP^M_�m ��_PZXg�]v^Q^QZ��PZks�[\V=Z�g�Zre3s T6R"YHTQU=ZX[os T�ZXe)YH]3R��=_O]3VX]3f3l

<PTos���TQTMe2s�_=T s �$])V�Z jdZ ePf�gL�PTa^QZXTQg�[6eOU)Z�gi49[ g [#�_d]=VXTQ4\gMZ YHZ�V [aR&s"] s _=[osd]3p=s-_PTEV [Ms"T!�*Zr]=^aT6e=T1�4? /� ��� 8�32��3��m

��_PT s�T Tos�_)[6R>T�giZ YGZ�Vr[QRPs"] s�_=]=giT�] p�" ?GF���%'���+/-��( 4'<=S3s=s _=T��P[aRq[ ^6]3eOT p"]=V�U)ZXgD�$TQ[�A=T6R&s _=[6e�S=gFS&[ V=Z e$s�_OT

V�Z jdZre&f�g+�PT6^ Z T g6ma��_PT�VXT6e=f3sX_�]3p&s�_dT YG]=V [aRFg��-��[3<PVXT �3#'['�!�PR"]3[Q^M_PTQg�s�_=T S��G�=TQR'V�ZkYGZrsd]3p&s�_DT�R-[Qe=f=T�p"]3R&s-_PZ�g

g��PTa^\ZrTQg 4��_PZ�VrT!s-_OT9ZrR5�$ZXUOsk_)]=p s�T6eHT�O^6TQTaU=g sX_dZXg�V�Z YGZ s>mQ��_PT4�=R>T6YH]=VX[6R>g [ R"T [\VXgM])Vr[QR-fPT6R>46s-_PT�p"]3R"T YG]vg s

s"TaT6sX_H���'J)[QeOU sX_PT��OTQR"gMZrg s T6e9s=���Ka#-<PTQZre=f�V�Tag g�R"T U9S=^ T6U=4\[�p-TQ[Ms-S=R>T��=R>])<=[�<PVkl�VXZ e�AOT6U;�$ZksX_)[#�P[QVX[os�T�V�T6g8g

e&[aR"R"]!�$TQU2[Me=s-TQR"ZX]3R>V ldm

1D]3Y5T ]9s�_OT R Vk[as"T ��V Z�]3^QTMePT �[ g sP�p"R"Z�^6[Qe5gL�OTQ^QZXYHToedg�YH[ol�[\VXgM]=<OT R>T6p"T6R"R"T9U�s"]#"$?+B4�+�� �%���E&m

��_PT6giT�ZreP^QVrS=UvT [�^6R-SPgF_DTaU2^aRq[aeDZnS&Y:pqR"]=Y+��[QTMs-]=V�Z)<PR"Z�TMpFVrl�U&T g ^6R>Z�<dT6U2[MePU;�=]3]=R"Vkl�Z�VXV SPg s�Rq[Ms"TaU8< l�A�S��6R>Z e

� K����vNDt �PV�m��&mrb\#8mQ�Ee)S=e!�vS�<PV�Zrg8_PT6U$g�A\SDVXVP� ,��0��/ � �-L�K\b N��=#'p�R>]3Y s-_PT I �!�dT6R&��]3YHT�A��$Z9�*TMYH<PTQR']3p

/.T g sP��SPR�A=[6e=[!_P[ g�T�G�=[6eOU3TQU�Vk[as-TQR-[\V!�=[ R�s"g�] p=s�_DT!e3SO^6_=[\V=^6R>Tag s��$Zks�_)[#�$T\V�V���YH[aR AOT6U$YBT U=Z [Me$e=]3s-^a_ �

s-_OT��PR-TQYH]=Vr[QR"g�[aRFTEVr[QR-fPTQ4\V�Z�A=T s _P]3giT�p�R>]3Y �$[QU=[6R8m\��_OT s�Rq[aePgMZks ZX]9e2s ]!�[QR"Uvg "$?GF!�&%����+/-��( Z�g�Z�VXVrS=g s�Rq[Ms"TaU=<=l

Page 18: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

gMVXZ f\_ s Vrll=]=S3e=fPTaR'g��OTQ^\Z YGT6e=gM4\g SP^M_)[\g�sX_PT#�=]3g s T6R"Z�]3R5�3[aR�sd]&p=s�_DTEg�AvSPV�V=��YG]��qbQL��-J=c��vc2pqR"]3Y 1O_=S3e=f3S=R"[

%7]=R-Y [as-ZX]\e2� T Y><PT6R&��J)���� �#>4'�_PZ�^M_2_P[Qg�Zks�g YH[3PZrYHS=Y ]v^9^ ZO�OZks�[\V!�$Z�U9s�_�[osPYGZ�U���_OT\Zkfv_3sPRq[Ms-_PTaR s�_P[Me

[6s=s-_PT4<=[\g8T�[\g�Zre "$?6F!��%����K/��O( m ��_PT�g�A\SPVXV9,�2� ��� 8�������2p"RF]3Y.s-_=T!,=�$1$��TQYH<=T6R7]3p&,�]P])<OZO%�]3Rq[EZXg

YG]3RFTEV�Z�A=T!s�_PTEVXZkjPZre=f$p"]3RqY:[QePUP4 [ Vks _=]\S&f\_)eO]9s'ZrU=TMe=s-ZX^M[\VX43^6[aeH<PTQgFs�<=TEZ�U3Tae3s"Zrp"ZXTQU2[ g,"$?6F!�&%����L/���(����$[aRFR"Z�gi4

K��3N��&4&K���� �3# m ��_PTQgiT�gKA9SdVXVXg [QR"TE^\Vr]&g8TEZke5g Zr`QT!s"] s�_OT YGT6[aeH]3p)" ?�F!�N%����K/���(a4)<vS3s'g8YH[QV�V TaR&s-_=[ae2sX_P]=g8T�p-R>]3Y

�$[\U3[aR m

%�T�� p-TQ[Ms-S=R>TQg�^6[Qe;<=TE]!<=g8TQR�jdT6U)]3es�_=T sXl��PTEg+�PT6^\ZrYHT6e�]3p +�4 '����� �-�I�� ��3���� �� BD�K�� �%��'E=4Q,��0�

,� � ���2p-R"] Y.,$[Qe=['�O]=Z�m ��_PT�U3]3R>g [QV!�PR"]3p"Z�VXT�Z�g Y5]3RFTE^6]3eO^ [MjPT!s�_=[6eHZre)[ V VPg+�PT6^ Z T g�]3p��4 )�+���������� ������! �4

[Qe=Us-_=Z�g�ZXg�^\VXTQ[6R>V l)[EU=T6R>Zkj T6U2p"Ta[6s�S&R"TE] p�"$��%) ����)(\m ��_OT�]=R"Z TMe=s�[as-ZX]\eH] p=s-_PT�]=^Q^ ZO�PZrs�[\V3[6R>T6[EZ�g

Z e3s T6R"YGT6U=ZX[os T4<DT s��$TaTMe=<P]9s�_HVXZkj Zre=f)gK�PTQ^QZXT6gMm\��_OT ]=R"Z�TMe=s�[as-ZX]\ePg�]3p=s-_PT#�&R ]9s-]=VX]��=_)[6ePUYHT6s�[\VX])�v_)[QR"T

gMZrYBZ�V [aR&s"]s�_P]vg TE]3p)"$?JB �+�� �%'��E2p-R>]3Y �$[QU=[aRF4'<=S=s=s-_=T�VX[�<PZX[QV��[QV�VXg!]3p s _PT s TQTosk_)[QR"T YGZ�g gMZre=f&ma,�� �

,� � ���)V [a^'A=g�[\VXV3s�_PT�[3�=]=YH]3RM�3_PZX^ p"T6[Ms-S=R>TQg�]=p s _PT �4 '�+���������� '�J�9�! +^QV [aU=T9m ��_=T�YH]=g sP^a] Y>�PV TMs"T�g�AvSPV�V

SPg TaU <=l��]=]=Z �MT6Rd[Qe=U � [6sXs-TaRFgi]3eH� K��3N\J&#'ZXe2sX_OT9ZXR'U=T6p"ZXe=Zks Z�]9e�] p M�#?�� B ���� �%���EH�[Qg�,��0��,� ��L�K 4 p"R>]3Y

�AP]3Rq[ 4Q[EV�]=^Q[QV�Z s�l;�=R>])<P[3<PVkl$l=]=S3e=fOT6R&s _=[6e2,$[ae=[��=]PZ)<3S=s=eO]9s-�=R>Ta^QZXg TQVrl�U3[os�T6U)�&��To_PR-TaePg YHTolOTQR"4 K��3N��3#Fm

�!RqSOg _PZre=f2_=[ g�[Q^\^aT6e3sXSP[os TaU s�_=T�^ ]9e=^Q[MjdZ s�l�]3p&s _PT�g�A\SPVXV!�&R"]=p-Z�V T\4Q[QePU)Zrs g R>T6gMTMYH<dVr[6eP^aT s ]�"$?GF��9%��!��/���(\m

��]=]=Z �8TQRD[aePU �'[Ms�s TQR"g ]\eH� K��3N3J't J��3#�^a]3R"RFTQ^Ms"Vkl$eO]9s T6U s _3[6s=s _PT s TaTos�_)[6R>T�^\]\YH�=[QR-['<PVrT s ] s�_O]=g T�]3p

"$?�F!�N%����K/��O( m ��_PTol$[\U3UPTaU s�_=[osPsX_dTol�U=Z p-p"T6R <3ls-_PT#�$T6[�AOTQR-�=[QR-[\^Q]\ePT p ]&VXUP4 <vS s=s�_PZrg�Z g�eP]9sP[!jP[QV�ZXU

U&Zrp"p-TQR"TQe=^QT\4Q[\g�s�_PZ g�p"]=V�U$Y [MljP[aePZ�g _�Zke�YH]3UPT6R"[os T��$TM[QR'ZXe2s�_DTEVXZkjPZre=f$p"]3RqY m ��_=T [ e3s-TaRFZX]=R-�=RFTMYH]PVr[QR"g

[QR"T�g YH[ V�VX49[ g�Z e "$?GF��9%��G�L/���(\m ��_dZrg�g�A\SdVXV&ZXg@<PTQg sdZrU=TMe=s�Z�p-Z�TQU2[\g-"$?GFC��%�����/���(\ma�-sdZ�g�S=eDVXZ9AvT\V ls"]=<OT�]&VrUPT6R

s-_=[ae2sX_dT�Vr[6s�T!� V ZX]=^QTMe=T3m � s-_OT6R'R"TQY [ Z ePg�U=T g ^QR"Z�<PTQU8<3l2��]v]&Z �iT6R'[ae=U �7[Ms�s TQR"g ]9e5� K��3N3J=#D[QR"T�p"R-[QfvYBTae s�[QR�l

[Qe=U�U3]eP]9sP[\U3U s ] sk_DT#A9eO]C�$V TQU3f=T�]=p s _PZ�g�gL�dTa^QZXT6g6mM��]��TojOTQR"4 s�_dTol�g _=]!�;s _=[os g8]3YHT s TaT6s�_��"V�Z�APT s�_P]3giT

pqR"]=Y+�$[ U3[6R"#5�$T R"TEgMVXZ f3_=s Vkl$YG]3R>T _3l6�Pg8]=U3]9e=s=s�_=[Me2s-_O]=g T�]=p)" ?GFC��%'���+/���(\m

��_PT YH[os T6R>Zr[\V=pqR"]=Y+��]3sX_P[6fv[QY _PT6R"T�R"TQp-TQRqR>TQU s ]�"$? B4�+�� '%��!E�Z eP^QVrS=UvT\g�g8T6jOT R-[ V=g�A=SPVXV=TQVXT6YHT6e3s�g6t

s-_OT YH]Pg s ^Q]3YH�PVXTos"T�[6R>T ,�� � � � ��J �vb)���2[MePU,�� �������-J����3NDK�p-R"]=Y+s-_PT�� �P[�A�*TaYH<PT6R8mQ��_PTMl$[QRFT

Page 19: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

<P]9s�_2R>T6p"T6R"RFT U s"]�"$?GF!�N%����+/���(4<=l�$[aRFR"Z�g�[6edU � T6['A=TMl��qJ=c=c��3# m ��_PTol�U=Z�g��PVr[6ls-_PT "$�&%� ��J�!(��-V�Z�A3T

YG]3R �v_P]3VX]3f9l�]&p=s-_PTEg�AvSPV�VX4)�$Z s-_�[�g s�R>]9e=f$U=]3R>g [\V=^a]3e=^M[6j ZksXlP4\[6edU2[!jPTQR�s"ZX^Q[ V=]=^6^\Z��=S3s��$Zrs�_s-_OT evS=^6_=[ V

^QR"TQg sPeP]9s T�vs"TMe=U=Z ePf;<=TM_DZ ePUsX_DT�^a]3e=U9l V T\gMma� _OT s�T6Tos�_)[aR>T�<=R"[Q^Q_3lO]=U3]\e3s-4��$Z s-_)[#�=[QR-[\^ ]3e=T p"]=V�U=��_PZ�^o_5Z g

�$TQ[�AOT RPs-_=[MeHZre "$?GF��9%��!��/���(a4Q[QeOU2[6R>T�gMZrYGZ�Vr[QRPs"] s�_P]3g8TE]3p!"$?JB �+�� '%���E��-Z eO^9VnSPU=Zke=fs�_=T!s�lG�PTQ#�p-R"]3Y

,$[Qe=[3�P]=Z3[QeOU �$[\U3[aR mM��]P]=Z �iTQRD[aePU �'[Ms�s TQR"g ]\eH� K��3N3J=#D_P[aU=�=R>TojPZ ]3SPg Vkl$R>T��O] R�s�T6U s�_OT�g��OT\^QZXT6g�pqR"]=Y

��]9sX_P[6fv[QY ]3e2s�_PT4<=[\giZ g�]3pP[gMZ e=fOVXT SG���PT6R'YG]=Vk[ R"4 ,��0� � �4� � �P4\]9eH�$_=ZX^6_$s�_OTol$ed]9s-ZX^QTaU s�_=TEV [ ^�A�] p

�&[aR"[Q^a]3e=T�p"]=VXU&m

�$�81P��IH1D13�>� �

����������� ��������������������������� ��� � �'* �� ����* ) ��,)���)� ����, � ��� ��2 � � �:��,:���� ����� ���

��]3RM�3_P]=VX]3fOZ ^a[\V=U=Z p"p-TQR"T6eP^aT\g@<=Tos �$TQToe ��?'(��� ;�! .[ae=U4"$?�F!�N%����K/��O(�[aRFT�YB]vg s Vkl$R>TQV [Ms"TQUs-]sX_DTaZ R

U&Z T s gM4 [ g!T6�DVr[\ZnePT6U8<3l���TaS3ePT6R'� K����9L=#8m ��_PT!_PT6[QU$]3p5�4?)( �9 ;�! �4��_PZ ^M_)p"TQT U=g�]\e)f=R-[\g g8TQgi43ZXg�_OTQVXU2[osd[ae

[Qe=f=VXT9mQ��sD_=[ g�[VX]3e&f�g�AvSPV�VX49[ae�]=^Q^QZ��3S3sdgFs-R"]3e=f=Vrl�Z eO^QV�Zre=TaU=�P]vg s TQR"]��-U3]=R"g [QV�V l;�$Zrs-_)[�e=SO^6_=[\V=^6R>T6g s

T�3s ToeOU&Zne&f;<=Tol ]9eOU s�_=T�^ ]9e=U3lOV�TQgM4Q[�f=Tae3s"V l�^\]9e=^Q[MjPTEU3]3R>g [\V!�=R"]3p"Z�V T\4Q[aeDU2[�eOToSPR"]=^6R"[6ePZrS=Y ]\eDV l�gMVXZ fv_ s Vrl

Z eO^9VXZre=T U2R"T\Vk[as-ZrjOT s"]s�_OT�p�[\^ TOmK"$�N%� '�J�)(�FC� %'�!�K/��9(a4'�_dZr^6_HZXg�[#<=R>]!�giT R"4\_=]=VXU=g�s�_OT _PT6[ U2YH]3R>T

_P]3R"Z ` ]3e3s-[ V�V lP4Q[QePUsX_PT�<=[ g Z�^6Rq[MePZX[QV=[ae=fPVrT s T6e=Uvg s ])^QVX]vgMT�gFS��=T6R>ZX]3RFV l�g SP^M_2s�_=[6s=s-_OT eOT6S=R"]=^6R"[6ePZ SPY Z�g

s"ZXVks�T6US!���[aRFU2R"T\Vr[6s Zkj T s ] s�_PT p-[ ^QT3m ��_PT�T=s T6e3sP]=p s Z�Vks Z�e=f)Z�g�e=]3s 49_=]!��T6jPT6R>46s"] s _=T�U3TQf3R>TaT�gMTaToeHZre2s _=T

fPToevSDg#� ����/6��%� ����!( 4'�_=TQR"T!s�_PT�]=^a^\Z��=S3sdZ�g�ZreP^6V�ZkePTaU$T jOT6e�YH] R T [Qe3s T6R>ZX] R"VrlPm313s SPU=Z Tag�] p=s-_PTE]9e=s�] fvTMe l�]3p

"$�&%� ����!( � ��T6Sve=TQR"4 K����3L=#'g s�RF]\e&fOVkl�g S=f=fvTQgFs=s-_=[6sdZrs g�g�A\SPV�V=g8_=[��OT�Z�g�U=T R"ZrjOTQU&m���T6S3eOT R ])<dg TQR�j TQU s�_=[os

U3S=R"Z e=f$U=TojOT\VX])�=YHTae3s=s-_=T�^Q]\eP^Q[oj Zks�l�]3p=s-_PT�S��!�OT R g�A=S=V�V!�=RF]3p"ZXV�T�Z eO^6R>T6[\g8TQg 4vZ�mnT9mn4Q_dZ g�[Qe=fOV T4/HU=T ^6RFTa[Qg8TQg

� %�Z f&m �v#FmvA R"]!�s�_)]3pPs�_dT evS=^6_=[ V=^6RFTag sP[aePU _O]3Rqe;<=]Pg8g Tag�p-[QZ�V9s"])p�SdVXVkl�T��PVX[QZXe2sX_dZrgM4'<=T\^6[aS=g8T s _PZXg

U=T6^aRFT6[ g TEg s�[QR�s"g�TQ[6R>VXZ�T6R'ZXe)]\e s ] fvTMe l��&<dT6p"]3R>T ��TMS3ePT6R'g s [6fOT! �=c)^6Y � ��T6SvePT6R>4dK����\L't L=L=#8mo��T�[\V�gi]

g _O]!�$TQU sX_P[os=s _=TE]3^Q^\Z��PZ s ]�� �=[ Vk[as�[ V3[6ePf=VXT�BD�HUvTQ^6R>T6[ g Tag!U9S=R>Z ePf s�_PT�Ta[QR"Vkl�gFs-[afPT���" �=c)^6YH#8mQ��_=SOg 4�<=]9s�_

s-_OT�]=^a^QZ��PZks-[ V3[ae=U=�=[QR"Z�Tos-[ V!�=R>]3p"Z V�TQg�[QR"T�V�TQg g�ZreP^6V�ZkePTaU2p>]3R �[aR>U)��YG] R"T1� ��L�,�����I�� ������! >�qV�Z�APT6#'U3S=R"Z e=fs�_=T

Page 20: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

TQ[6R>V l�g s�[ f=TQg!��" �=c)^MYH#&s-_=[ eHZ e2s�_PT [ U9SPVrs"4'�_PZX^o_5Z�g�^QToR-s�[\ZreDV l$SPePVXZ9AvT sX_dT [Me=^\Tag s�Rq[ V Y5] R��=_O]9s�lG�PT�]3p <d]9s�_

V�Z jdZre&f�g+�PT6^ Z T g6m

�.V�Z�APTQVrl$[6eP^\Tag s�] R'p ] R-<P]3sX_HV�Z jdZ e&f�g+�PT6^ Z T g!ZXg �� ��L�,�����I�� '�3���! 0/� ��! ��2 �3�P�-]=p s�T6eHZre=^Q]3R"R"T\^os"V l

^Q[QV�VrTQU�"$��%' '�J�)(CB ��%3��21%�/��������)( � gMTQT�AET6R"[6[\U=gM4&K������\#>4 �_dZr^6_5Z g�YH]=R��=_=]=VX]3fPZr^Q[ V�V l$[QePU$TQ^Q]3VX]3fOZ ^a[\VXVkl

Z e3s T6R"YGT6U=ZX[os T4<DT s��$TaTMe2s�_=TQY m���TaS3e=TQR'� K����3L=# g _=]!�T U�s�_P[osPYG]=g sd]=pdZks g�^QR-[QePZX[QV3[ e=fPVrT\g [aRFT�[QV�g8]

Z e3s T6R"YGT6U=ZX[os T3m�� '������������ ������! 7/� ��� �32��3�PZ g �$T\VXV!A3e=]��2e�Zre2s-_OT�TQ[ag s T6R"e2� TaU&Z s T6R"R-[QePTQ[MeHV [Ms"T!��ZX]v^QToePT 4

T\gL�OTQ^QZX[QV�Vkl�p-RF]3Y �7Z�A=T6R"YGZX4 1=[aYG]&gi4 �7TMe3s-[\V ]!�3_=]=gi4Q[Qe=U � [6R"[af=_PTo_H�-��_PTae=ZrSPgM4&K��&b3b��vATaRq[a[QUPg 4&K��������

ATQR-[Q[QU=g�[ae=U ,�]3S=p"]=gi4 K����=c=4Q[Qe=U2R>T6p"T6R>T6eO^9Tag�s-_=TQR"TQZne&#FmM $T V [Ms"TaU2p"]3RqY5g!ZreP^QVrS=U3TEV [Ms"T�YBZXUvU=V T!�*ZX]3^ T6e=T

�pqR"Z�^6[Qe)R"T'�=R>T6giToe3s [os�Zkj T6gM4\g SP^M_)[\g�� �6��� �-��%� '����(�p-R"]3Y %�] R�s �!T6R-e=[Med4 ,�Tae lv[\4Q[aeDU=�OT6R-_=['�Og �ETaePZ9��T V�V [ V�4

� ]3RF]v^Q^6]�� ��]=]=Z �MTaRF4&K��������vA�S��6R>Zked4&K��=N��3# m�&"$�&%� ��J��������/���( � BD�3�� �� ����!(���RF[aY$<=]3S=R-fP4&K��&b �=43p�RF]3Y s�_dT

TQ[6R>V l�V [Ms"T �CZX]&^aT6e=T�]3p7�VrfPT6R"Z [��q��R"[QY$<=]\S&R-fP4&K��&b ���vAT6Rq[a[ U&gi4 K������3#'^Q]\SPV U8<PT s�_OT�TQ[6R>VXZ�Tag sPYGT6YH<PT6R']=p

s-_PZXg�f3R>]9S��DmK"$�&%' ����!(����!(����� �&� (EA�S��6R>ZreP4QJPc3c3cP49p-R>]3Y s-_OT YGZXU3U=V�T��CZr]P^6T6eOT�]3p �$[aYGZ9<PZX[�ZXg�eP]9sdZreP^QV SPU=TaU

<PT6^a[QS=giT�Z s'ZXg <&[agMTaU+SG�=]3e)[!s�_dZkR"U$YBTMs-[a^Q[6RM�=[ V�m

�"e$�4? /� '�! 1��2 �3��4 s�_PT�e3S=^a_=[QVP^6R>T6g sP[aePU2^a]9ePU9lOV�TQg�[aRFT�[��G�&R"]!PZ YH[6s T\Vkl$[Ms=s-_PTEg [6Y5T�V�T jdTQV�4

[\V s-_O]3S=f9_$s�_PZ�g [aR>T6[EZ�g�]3p�s"TMeH^6R"S=g _OTQU)U\S=RFZ ePf)p ]&g8gMZXV�Zr`Q[Ms Z�]9eP4��_PZX^o_H�=R"TMjdToe\s"g@�PR"TQ^QZ�g8TET6g s�ZkYH[6s TQgom �ePT ]3p

s-_OT�<PTag s��=R"TQg T6R-jPTaU)g�A\SPV�VXg���p"R"]=Y 1P[6YG]vgM4 ��8�E#'_=[ g s _=T�e3SO^6_=[ V&^6R"T g sd^QV�T6[QR"Vkl$YG]3R>T��O]3g s"TaR"Z ]3RDs�_P[Me2s-_OT

^Q]9eOUOlOV�TQgoma��s7g8_=]3SPVXU=<=T!eP]9s�T U s-_=[Ms'A�[aS=U=R�lPW g�U=R-['�$Zre&f��-A�[QSPU3R-lP4DK �)�3J�� K �)�=NDt �PV m J=NP49p"ZrfDm JP#'g8_D]!�$Zre=fs�_OT

RFTMjdT6R"g T�RFTQVX[os�ZX]3e=gF_DZ �HZre)[Eg�A3SPV�V3p-R"]=Y+� Z�A=TQR-YGZP� � ��$�� �vg�A=SPV�V "���ZreHATQR-[Q[ U=gM4dK������\#7ZXg�ZXe=^Q]3R"R"T\^ s �

s-_OT e=S=^M_P[QV=^QR-T\gFsdZ�g�Zke�p-[Q^os VXT6jPTQV��$Z s�_$s-_=TE^6]3eOU9lPVXTQg�Z e2sk_PZ�g�g��=T ^QZ YHTMeDm��4 ������������� ��3���� /� ��! 8�32-�3�PZ�g

[\VXgM])Z e=s�TQR-YGTQU=Z [Ms T�Z eH<P[agMZr]=^Q^ ZO�OZks [QV3YG]=R&�3_P]=VX]3f\lP4'�$Zks _)[#�)T6[�As-S!<PTaRF^QVXT�[6eOU2]!�OT6eH<=[ gMZr]=^6^\Z��PZrs ]��

g��v_PT6e=]=ZXU2[MePf=VXT9m ��_=T�g [aYHTEZ�g�s�R"S=TE] p=s-_PT�[ojdTaRq[af=T#�=]3gMZ s�Z ]\eH]3p=s-_PT [ e9s"TaRFZr]=R']3RM<PZks�[\V�<P]3R"U=T6RD[QeOU)]3p&s�_DT

Z e=p-R-[\]3RM<PZks�[\V=p-]=R-[QYGT6eH��%7ZXf&m�� #Fm ��_OT��O]=g Zks ZX]9e�]=p s _PT e3[ g [\V3eO]Os�^a_2j=[aRFZ�TQg�V�Z s�s�VrT [QYG]\e=f s [P[ 4)<=S\sdZksPYH[6l

<PT�giV�Zrf3_3s�V l$YG]=R"T [Qe\s"T6R>Zr]3R7Zre "$?�F!�N%��G��/���(�s�_=[MeHZXe)]9s�_=TQR'gK�PT6^\ZrT\gMm

Page 21: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

��_PT U=ZXp-p"T6R>T6eD^aT�ZreHgKA3SPV�V=g _P[3�=T#<PTos���T6Tae2s�_PT sN�])V�Zkj Z e=f�gL�PTa^\ZrT\g�ZXg�R"Tap"V�TQ^os TQU�Z e2s-_OT\ZkR g�A9SdVXV

YGT [ gFS&R"TaYBTMe3s g��"A�S��QR"ZXeP4dK����\c=# �vZre2% ZkfvS&R"T!J "$?6F!�&%����+/��O( �"U3[Qg8_=TQU)VXZXe=T��$Zrs�_)[QgFs T6RFZ�g�AOg #'Z�g�^6]=YH�=[aRFTaU s ]

�4?)(3�� ;�! :��SPg T U2[Qg�[Eg s�[Qe=U3[QR U #Fm � _OT [6j T6R"[6fPT�YHT6[ g S3R>T6YGT6e3s g�]3p s�T6eHg�A\SPV�VXg�]3p-�#? /� ��! ��2-�J�&p-R>]3Y s�_PT

�[\g s T6R"e2� TaU=Zks T6R"R-[QePT6[aeHVr[6s T � ZX]=^QToeOT�[6R"T �PV ]OsXs T6U�]9es�_dT�g8[QYHT�U=Z [af3R"[aY ma�*]=gFsd]3p&s-_PT6Y p-[ V�V!<PT6sN�$TQTMe

s-_O]=g T�]3p&s�_DTEVXZkjPZre=f�g��PTa^\ZrTQgom ��_=TEV�Tag g �=]=g s T6R>ZX] R"Vrl�T��3[aePU3TaU2e=SO^o_P[QVP^6R"TQgFs'ZXg�RFTap"V�TQ^os TaU)Z e$s�_OT�VX]!�

jP[QVXS=TE]3p����3#7^Q]3Y>�&[aRFTaU s ]H�4?'(��� 8�� �4 [aedU)Z e2s�_PT�g8_=]=R s��P]=g s�]3R�<PZrs-[ V!�P]3R�s ZX]9e5]3p&s�_DTEg�A3SPV�V=� �3#Fma��]!�$TojPTaRF4

s-_OT��PR-])�P]3R-s Z�]\e)]=p s�_dT [ae s TQ]3R <PZ s [\V3^a] Y>�=[6R>T6Us"] s�_=T#�=]=gFs-]3RM<PZks�[\V��=]=R�s"ZX]3eH��� ��� jdgom�� ���-#7Z�g�YG]3RFTEg Z YGZ�Vk[ R

s"]H�4?'(��� 8�� ;s _=[6e$s�]�" �9%� '�K�!(a4)�_PZX^M_)_=[ g [Eg _P]3R-s T6e=TaU2YH[� Z�VXV [aR�l�R>T6f=Z�]9e'm'�4 ������������� ������! 0/� ��C 1��2-���PZ�g

[\VXgM]2YH]3R>T�VXZ9AvT1��?�(3�� 8�! :ZXe2_=[MjdZre&f$[MeH]3^ ^QZ��PZks�[\V9eO]OsPYHSO^6_H<=R"]3[aUPT6RD[6s=s _=T�<=[ g T!s�_=[6e)[Ms=s-_OT e=SO^6_=[QV

^QR"TQg s>mQ��_PT#�=V�T6giZ ]3Y5] R��=_OZ�^�gqs [os�T ]3pd[U=TaT��)e=[ g [QV3eO]9s-^M_H�"VX]3e=f2e=[ g [\V!<O]3e=Tag #'UPZ g s Zre=f3SPZXgF_=T\g �4?'/- ��! ��2��3�

pqR"]=Y <d]9s�_HVXZkj Zke=f�g��OT6^\ZrT\gMm

�� ��L�,�����I�� ������! 7/� ��! ��2-�J� �[ g@�&R"]!<=[3<dV l$[�YBZ�PT\U2p T9T U3TQR-<PTQ^6[aS=giT�ZrsdZ�g�YH]3R �v_P]=VX]3f3Z�^6[\VXVkl$[aePU

YGT s�R>ZX^ [ V�V l�ZXe3s-TaRqYGTaU&Zr[6s�T4<dTos���T6TaeH<=]Qs�_5V�Z jdZre=f)gK�=TQ^QZXTag6mM�'RFTMjdZ ]3SOg [ gMg ZXf\e&YHTMe3sd]3p&s _PZ�g�gL�PTa^\ZrT\g s ]

"$�&%� ����!( �-T3m fDm 4=A�S��6R>ZreP4dK������P4QJ=c=c3c��Q��]=]=Z �iT6R"4 K��3N��\#5�[\g�YBZ�g V�T6[aU&Zne&f8<OTQ^6[aSPg TEZksdZXY �PV�ZrTQU)Z s��[Qg�[

<&R"]!��g T6R8m

7 �#7 �<�<, � ��� � 2���� ���� ��+� ��� � ,1��� / ��� � , �1���

��_PT6RFT�[6R>T�]\ePV l$[p"T��.V [Ms"T!� Z�]3^ ToedT �p-R>ZX^ [aeHgMZ s T6g��$Z s _HVr[QR-fPT YH[aYHY [ V g6m ��_PTEg ]=VXT�RFT3�=]=R�s']3p

�4?)/� ��C 1�32��3�PZ�g�[�gMZrePfvV�T s ]=]9s�_�p-RF]3Y 1=[a_P[3<PZ�46��Z�<3lO[��&�$TaR-e=]3R'T6sP[\V�mn4&K����\N3#>4'<&S3sdgi]3YHT�]Osk_PT Rd�pqR"Z�^6[Qe

��Z�^aTQR"]Os ZrePZ3[QR"T]3pd^Q] Y>�=[6R"[�<dVrT�Y�] R��=_O]=VX]3f3l&m�&"$��%� '����( � ���G�!���J�9 �/�(3��(�A�S �QR"Z eP4 K������=4Qp"RF]3Y?� SvePZ giZ [

�qA�S��QR"ZreP4&K������ � AT6Rq[a[\U=gM4&K���� �3# ^Q]9SdVrU8<PT�^a]9edgL�OTQ^QZXp"Z ^#�$Zks _H�4?'/� '�! � 2-��� 4'<=S3s=s�_PT�]3ePVrl�g��=T ^QZ YHTMe

U=T6p"ZrePZks�TQVkl$R"T\p-TQR-['<PV�T!s�] s�_PZ�g�g��OT6^\ZrT\g�ZXg [Eg�A=SPVXVP� %�13� � K��3N3L �3#&s"]v])ZreP^Q]9YH�PV�T s�T p"]3R'Zrs"g �=RFT ^QZXg T [ pqp Z ePZks Z�T6g

s"]8<PT [\g8giTQg g8TaU m ��e)Sve��3S!<dVXZXg _=TaU2[QeOU=�=]=]3R>V l;�&R"T g T6R-jPTQU$Y [3PZ�V�Vr[�p-R"]=Y+� ZXgMg [\g8p-[Q4Q�*]3R>]3^Q^a]P49U3[as�TQU2[Ms

Page 22: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�3�

^\Vr]&g8T!s-]sX_DT!�*Zr]=^aTMePT'� ��V�Zr]=^QTMePT4<=]\S=e=U3[QR�l5� E[6lOe=[\V=Tosd[QV-m 4�K������3#"43ZXg �=R"ZXYBZrs ZkjPTEZre2s�_PT _PZXf\_H^Q]9evePT6^Ms"ZX]9e

] p=s�_DTEVXZke=fvS=[QV3s�S�<=T6R>^QVXTag!]3p � J�m

�D��] s-[�O[E^6[6e><=T�ZXU=T6e\s Z p"ZXT\U)Z e2s _=T 1=�GKEgFS�<=YHT6YH<PT6R"49��]!��TQR��1�[Qg _$<$[QV�V�Tol��"^QZ R"^Q[ �&m � ��[Q#Fm

��ePT�]3p=s-_PTQYG4���?' 8�����J�����!/6�&%��C �4Q_=[ g s�T6Tos�_Hg8_=]!�$Zre=f$gMTojOTQR-[\V=]3p&s�_dT [��=]=YH]3R��3_PZ�^ p"T6[Ms-S=R>T6g�]=p

�4?)(3�� ;�! �4Q[Qe=U s�_vSPg sX_PT6R>T�ZXg�V�ZksXs�V TEU3]\S�<3s=s�_=[as=s-_=TMl;<PT6V�]9e=fs-]s-_=TEg [QY5T�V�ZneDT6[af=T\ma� _PT�]9s�_=TQRD_=[ g [

U=Toe3s�Zks ZX]9e)[ae=U�g�A3SPV�V3YG]3R��3_=]PVr]=f9l$R>TQgMT6YH<PV�Zre=f�"$?GF���%'�!�+/���( 4 p"]3R5�_dZX^6_HZ s'ZXg�[EV�Z9AOTQVkl$[ eP^6TQgFs ]3R mM��TQg��PZ s T

s-_OT S=e=[aYH<DZrfvS=]9SPg@�v_3lPVX]=f=TMePTos Z ^#�=]=gMZ s�ZX]9eH]3p=s-_PZ�g�giT6^a]3e=U)gK�=Ta^\ZrT\g 4"$��%' ����)(CBD�K�� '%���EP4\ZksP_=[\g@<OTQToe$s-_=T

g SG<��MTa^ s ]3pDYHSP^o_5^Q]9e=p-SPgMZr]3eDma�"e2s-_OT�V�Z s�T6R"[os�S=R>TQ43Z s��$[Qg!^a[\VXV T U)TQZksk_PT6R!"$?�F!�N%����K/���(���[6s=��]3sX_P[6fv[QYG46�$[\U3[6R>4

[Qe=U;�=TQR-_&[3�Og /0TQgFsP��S=R�A=[6eP[6# ] R5�4 '�K���������� ��3� �C B �+�� �%���E)��[6s=,$[ e=[��d]3Z3[QeOU=�=TQR�_&[��=g ,�]v])<dZ9%']3R"[a# m ��_OT

p"]3RqYGT6R7Z�U=T6e9s�Zkp"ZX^Q[Ms"ZX]3eHZrf3eO]3RFTQg�sX_PT YG]3R&�=_O]=VX]3fPZr^Q[QV=UPZrp-p"T6RFTaeP^QTQg@<PTos �TQTMe2s�_=T!s��$])gK�=Ta^9Z TQg�]3pG"$�9%� '�K�!(o4

<=S3sdZ�g@�v_ lOV�TMs"Zr^Q[ V�V l�gMToedg Z�<PVXT9ma��_=TEV [Ms�s TQR"43]3R"ZXf=Z e=[ V Vrl$SPgMT6U8<=l��]=]=Z �iT6RD[QeOU �'[6sXs T6R>g ]3eH� K��vN9J=#>4Q[\g g8S=YBTag

s-_=[6sdZ s ZXg sX_dT [MeP^QTag s�]=R']3pDVX[os�T6R �4 '�K���������� ��3� �C HmM��]!�$T jPTQR"4 s T\Tos�_Hg ZXYGZrV [aRDs-]s-_=]Pg8T�]3p&s�_DT!s l6�PT�]3p

K�4? ��B �+�� '%���E2p-R>]3Y ,$[6eP[3�P]=Z=U=ZXg��PV [Ml$eP]3e=T�]3pDs�_DT�[3�=]=YH]3RM�3_=Z�^ p"T6[Ms-S=R>Tag�]=p-�4 '�K������� �6 '�����! �4Q[QePU2[ R"T

[\g gi]3^QZX[os TaU=�$Zks _ "$��%� ����!( g�A9SOV�VXg6ma��_=T6R>T�Z�g�VXZrs�s VXT�U3]3SG<9sPs�_=[6s=s-_OT l;<PTaVX]9ePf s ] s�_OTD"$�&%� ��K�!(�^QVX[aU3T3m

� s�s"R"Z9<3S3s�ZX]9eH]3p&s�_PT�YHZXUvU&VrT!� V ZX]=^QTMe=T "$? B �K�� '%��'Es"]H� ������������� '�J�9�! �[Qg�s-_OT�g s�[QR�s"Zre=f;�=]PZ e=sP]3p&s�_PT

^Q]9e=p-SPgMZ ]3eP4 �_dZr^6_>�[Qg�p-S=R-s�_PT6R'ZXeP^ R TQ[QgiT6U=<3ls _=T�[\g gMZ f=e=Y5Toe=sP]3pDs�_DTEVr[6s�T � VXZX]=^QTMe=T s RqSOT � ������������� '�J�9�!

s"] sk_PT�g [ YBTEg+�PT6^QZXTQg6mM ET6j Z giTQU)ZrU=TMe3s"ZXp-Z�^6[6s Z�]\ePg�]3p=s-_PT!YG]=g sdUPZr[Qf\e=]=g s�Zr^g+�=T\^aZ YGToePg�[aRFT�f=Zkj Toe5Zke���['<PVXT

L�m

� jP[ Z�Vr['<PV�TETojPZXUvT6e=^QTEgFS=f=f=TQgFs-g�s�_&[Ms=s�_=T sN�$])V�Z ePT6[QfOT6g!g+�PV�Zksdg8]=]9e)[ap�s"TaR&s-_PT!�*ZX]3^aT6ePT�� �7V�ZX]v^6T6eOT

<P]9SvePU3[6R-lO4=V TQ[ U=Zre=f2p"R"] Y:[Qe)[ae=^QT g s�R"[ V YGZ�dT6U��-p-T\TQU=TaRF46s ]2[VXZXePT [afvT�] pDfvRq[a`vT6R>g����4 ��+�5�����I�� ������! $#'[6ePU$[

V�ZrePT6[QfvTE]3p-<=R"]C�$g8TQR"g��O"$�N%' ��J�!(a#Fm\��_OT�Z eO^QR"T [ag8TQU$[QU=[3�3s�[os�Z�]9e�]3pDs�_PT8� ��K������� �6 '��� �C 0^\Vk[ U=T s ]$])�dToe

f=R-[\g giV [QeOU3g�[Qe=U�g [ojO[6eve=[\g�VXTaU s-_OTD"$�N%� ����!( ^QVr[\U3T!s ])gF_DZrp�sdZks g!T ^6]PVr]=f=ZX^6[ V eOZ�^M_=T s�]��[6R>U=g�YH]3RFT�^QVX]=giT6U

_&[3<PZrs-[Ms gom � _PZXg �[Qg�[\gig ]3^ Z [Ms"TaU=�$Zks�_H^M_=[QR-[\^os T6R7U=ZXg+�PV [a^QTaYHT6e3sFm\1PT6j T6Rq[ V=p-TQ[os S=R"T\g�]3pC"$��%� �����( TojP]vVkjPTaU)Zre

U&ZrR>T6^6s ZX]9eOg!])�G�P]3g Zrs T s�] s _=]=g8T�p"]=V�V ]C�$TaU <3l �4 '����� ���I�- ��3���� GtQg8Y [ V V�T6R7g ZX`QT\4\g s RF]\e=f=T6R-�P[aRq[ ^Q]3e=T�p"]=V�U=4

Page 23: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�3�

YG]3RFT!s-R-[QeOg jPT6R>g8T�VX])�=_Pgi4 s�RFZX[6e=fvSPV [aR&� �=4\[6ePU$_=]=VXU3Zre=fs-_=T�_PT6[aU)V�T6gig�ZXe=^QV�ZkeOT6UP4'�$Zks�_2s�_=T�[\g gi]3^QZX[6s�TQU

p"T6[6s�SPR"T9g�]3pP[Qe3s"T6R>]3Uv] R"g [ V�V l�ZXe=^\VXZkeOTQU)]v^6^\Z��dZ s-[ V�4QYG] R"T�^Q]\e=^Q[ojdTEg�A\SdVrVC�=R>]3p-Z�V�TQ4Q[QeOU)g s�RF]\e&fvT6R

<&[ g ZX]v^Q^QZ��PZks�[\Vv^ R"T g sP[6eOU s-SG<=TQR"^QVXT3mo�j ]=VrS3s-ZX]\e)pqR>]3Y YGZ�PTQU2p T\TQU&Zne&fs�]8<=RF]!�$gMZre=f�Z g�])�G�P]3giZks T s ]=�E_&[Ms

_&[ g�[QVXYH]=g sPS=ePZ jPTaRFg8[\VXVkl;<PTQTMe)Uv]&^MS=YHT6e3s TaU2p"]3RDSPe=f\SDV [as-T\g��-T3m f&mn4 � T R-e=j=[ V�V3[ e=U % ] R�s�TQV�ZrSOg 49JOc3c3J=#>4'<=S sO_=[\g

<PT6TMeHU=T6Y5]9ePg s R-[6s TQU)Zre�T9VXT3�3_P[6e\s"g��q��T6R>VXZre=f)T6sP[ V�mn4&K������3#>4Q[QePU8�=RF]�<&[3<PVrl�]=^Q^oS=R"R>T6U)ZXe "$��%� ����!(�[Qg ��TQV�V�m

���,��� /*����HA �w��$�!1

��s�_=[Qe�A$� s�] � [ R-[ �$[ Z�VXTQY)[QR"Z [ Y [QePU sX_PT���S3s-_O] R"ZrsXl)p"]3RD �TQg8TQ[6R>^o_�[Qe=U)��]9ePg TQR�jP[Ms"Zr]3e5]3p

��SdV s�S=R"[QV3��T6R>Z s-[afPT [ae=U s�_PT �p-[ R�R"TQf=Z�]9e=[\V=gFs-[Ms-T�p"]3R'[\V�V�]!�$Zke=fs�_=T!��Z9A=Z�A3[! �TQg TQ[aRF^M_2�'R>]��iT6^6s=s ])^a]9ePU\S=^6s

pFZXT\VXU=�]=R�A�Z e2s _=T � ]C��T6RD�1�2[\g8_P49[aePU /�sP� [ YBZ s S �Z�VkYH[Ep"]3R �=R"]3jPZ�U=Zre=f2p-[ ^QZ�V�Z s"Z�Tag�s"]=��]3R A�Zre2s-_OT

�$[6s Z�]\e&[ V9��SPg TMS&YG4Q�U=UPZrg�� <P[�<P[9m ��Rq[MjdTaV=T�G�PTMePg Tag�ZXe2�s�_PZX])�PZ [�[QeOU ,$TMe=l3[#�$T6R>T�p-S=e=U=TaU=<3ls _PT

��� 21'mM%7ZXTQVXU=�$]3RMA2[MsP��ZOAPZ�A=[#�$[Qg�g S!�G�P]3R�s TQU8<3ls-_PT��$[Ms"Z�]\e=[\V=ATQ]3fvRq[��=_PZr^21O]3^\ZrT6sXlP4\V�]3fOZ�g s�ZX^�gFS!���O]3R�s

�[\g �=RF]9j Z UvTQU8<3ls-_PT �"ePgFs"ZrsXS s T�]=p �$S=YH[QeH��R"ZXf=ZXePgM4 [aePU sX_dT�Zne jO[QVrS=[�<PV T�_=T\V���] p=s-_PT ��gMg [#�PTa]!�PV�T YH[aUvT

Zks��O]&g8gMZ�<dVrT3m\1��PT6^ Z [\V9s�_P[Me Avg�s"] ��ma�V�T6YGg T6fOTQU=4\A5mM�^3AP4Q[QePU �$mM,�Z Y$<=T\V3p"]3RD_=[6j Z e=f�T6e3s-R-SOg s T6U2YGT#�$ZksX_

s-_OT�g sXSPU9l�]3pDs�_DT!��]!��T Rd� �[ g _)R-_DZreP]vgom\��_=[ae�A=g�[\V�g ]�s"]�;ma��T6['AvT6l$[6ePU�m6� <3S=[E��,���#"49��m � [\g8g lO4

%�m6 �TMeP]9SPVrs"4Q[6ePU �6m � SOZ�g ZreH� ������'#�p ] Rdf=Zkj Z e=f$[Q^\^aT\g g�s ])^a]&VXVXT6^6s ZX]9edg�Zne2s�_PTQZrR�^a[6R>TQ49[aePU s ] $m�� ])<��\4

�Qm\1=S&R-[ SPVks 4 �Qmo/*lve&eP4Q[QePU$TQgK�=TQ^QZX[QV�V ls"] ��m\��V TQYGg8T6f=TaU2[QeOU �HmM �TQT6U$p"]3R-�D]=V�ZXg _=Zre=fs-_OT �ePf=V�Zrg _DmM� [Qe3l

SPg T6p-SPVP^a]3Y Y5T6e3s gD<3l �QmM� mM�$[\R"RFZXgM46�Bma�7R"]OsX_dT6R"]=4Q[QeOU2�)m��Qm6/0Z e�A=V�T6RDf=R"TQ[Ms"V l�Z Y$�=R>]9jdT6U s�_OT

YH[Qe3SPgi^6RFZ � s"m

Page 24: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

���

���>��� E���I !����"����

�V�T6YGgMT6fvTQU=4 ��m 49[aePU �GmQATaRq[a[\U=gom J=cvc=c&mQ�*eDT��*��ZXUvU&VrT!� VrT\Z g s ]=^6T6eOT p"[6S=eP[�p-RF]3Y s�_PT4��S=gMZrUPZrYH[��

��T\Vk[ Vk[�A2R"TQf=ZX]\eH]3p&s�_PT��p�[\R"4 �s�_DZ ]!�PZk[vmQ��]3YH�3s�T6g� ET6eOU9SPg�U3TEV�W ��^6[\U��6YGZ TEU3TQg!1P^QZXT6eO^6T\gi4

1P^\ZrT6eO^6T\g�U=T�VX[���TQR-RFTETos U3TQg ��Vk[ae��6s�T\g ��� KOt bvL����Fb3b��&m

�Rq[aY$<D]\S=RqfO4\� m=K���� �=ma� [ Y YGZrp��6R>TQg�p"]=gMg Z�V T9g�U\S2��[ R"]3^3mM� �6YH]PZrR"TQg�UPT�Vr[21O]=^\Z �6s ��U=TQg 1O^\ZXToeP^QTag

�$[os S=RFT V�VXT6g�U3S2��[QR"]=^!L��DtXK��qN3L'm

�Rq[aY$<D]\S=RqfO4\� m=K��3L��=m �!]9e3s R"Z9<3S3s ZX]9e��V�W �os�SOU&T�f��a]PVr] fvZ �9S=TET6s��=[\V � ]9e3s ]=VX]3fPZ �3S=TEU9S;<&[ g giZke5U9SHVX[a^

E]3U=]=V��v_=TTos U&TEVk[#<P[agMg8T j=[ V�V��QT�U=T�V�W ��YG] ���!�'m6J ��K�� b �9JHZreH��ma�R"[6Y$<d]9S=R"f)�-T\U&m #"4Q�*ZXgMg ZX]9e

g ^\ZXToe3s ZXp-Z �3SOT�U=TEV W ��YG]Dmo<�]=V-m�KvtaA��\]=V ]3fOZXT�Tosd��e3s�_=R>])�=]PVr]=f=ZXT9mM�UPZ s"ZX]9ePg�U9S$� SOg��6S=YG46�7[6R>ZXg6m

�Rq[aY$<D]\S=RqfO4\� m=K��&b �&ma<�T6R�s ��<=R � g�^a]3e3s"Z ePTMe3s-[MS��U9S$��Z�]3^��6e=TEg SG��� R"ZXTMS&R U=T�V�W �p-R>Z�� SOT�U9S$�H]3R U m

�'S�<PV�Z ^ [Ms"Z ]3e=g�U9SHgiT6R�j Z�^aT�U=T�Vr[^6[aR-s"T f �6]PVr]=f=Z��9SOT�U=TEV W ��V f��QR"Z�T���ed]9S3jPT V�VrTEg��6R>ZXT #Fma�'[ V �Q]3e3s ]=VX]3fvZ�T

L't K�� K\b��&m

�Rq[aY$<D]\S=RqfO4\� m=K��=N3c&ma��T6g�<�TQR�s��3<PR �9g�U\S2�7V �QZXg s ]3^��MePT�U=T�V�W �2p-R>Z�� SPT�U\S2��] R U ma�R>^6_PZkjOT\g�U9S$� SPg �oS=Y

�$[os�Z�]\e=[\V=U=W ��Z�g s ]=ZXR"T �$[os�S=R>TQV�VrT\4\g��QR"Z�T N=4DKacDt K�� KaJ �&m

�Rq[aY$<D]\S=RqfO4\� m 49[aePU �QmM�7Zkj T s T [aSDm&K��9J��&mM��TQg�<�TQR�s��3<PR �Qg�U\S2� ]3e3s"ZrT6e5U3T 1O[\Vr]3ePZ��9SPT3ma��e=eP[QVXTag�U=T

�'[\V��Q]\e3s-]=VX]3fPZrT2K �Pt K�� �\JDm

��T6_=R"ToePg Y5TolPT6R>49�$m6,HmDK��vN��&mo� ]Osk_&[af3[QY.�HZXV�V 4Q,$[6eP[��=]PZ 4\[6eOU �A=]3R"[=ta[�f=TaePT6Rq[ V=g S=YHYH[6R-l�] p

g s Rq[Ms"Zrf=R-['�=_3l$[QeOU)p-[6S=eP[ �����Dm=K������ KaN=c)Z e �)m\��])�G�PToePgi4 %�m ��mM��]��$TQV�VX4vA2mo��m\�"g [a[\^Q4\[QePU $mM�mM%�m

��TQ[�AvT6l��-T\U=g6m #>46�[QR"V�Z�TQg sP��[ae)[aePU �e3j ZkR>]9e=Y5T6e3s g�Zre2s�_=T ��[�A=T! �S=Uv]&Vrp �E[ gMZreDm6I$ePZkj TQR"g Z s�l�]=p

��_PZX^6[QfO] �'R>T6gMgi43��_PZX^6[Qfv]&m

��T R�eP]3RF46 $mM��m 4 ,Gmo��T Z�g gMZrfP4Q[QeOU �Hm ��]!<OZ�TMeDm&K���� N&m6�[MR>Vkl� VXZ�]=^QTMe=T!� T6RFZXgig ]=U3[a^6sXlPVr[EpqR"]=Y 1P[a_=['<PZ�4

��Z�<3l=[ ��� �DmaJ������-J&b LHZre2�Hm3�$]3[M`v4Q�)mM�V=�Rqe=[aS=s-Z�4Q�)mvA�[6`\ZXR�l 4 �QmQU=T!�HT Zke=` T\VrZreP49[aePU �Hm6�Hm3��]=[6`

Page 25: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

���

�qT U3g6m #F4Q��T9]3fvT6eOT �7[ V�Ta]9e=s�]=Vr]=f9l$[aeDU)ATa]=VX]3f3l�]3p�1P[a_=['<PZ�ma��V [ae2��Z�gigM4 ��T'� �$] R�ADm

�ERF]!�EeD46%�ma�Hmn43[6ePU���mv1DmM% TQZ�<=TQV-m K���� Kvm\13s-R-[6s ZXf3R"[��P_3lO4=U=T�P]vgMZ s�ZX]9e=[\V=TMe jOZXR"]9ePYHToe3s gM4Q[aeDU

�=[\Vr[QTa]=f=Ta] fvR-[��v_ l�]=p s _PT ,�]=]�<PZ9%7]3Rq[!%7]3R"YH[os�Z�]\e �'���Dm=K����=c)Zre �QmM� mo�$[\R"RFZ�g��-T\U&m #"4Q,�]=])<dZ9%7]3R"[

�TQg8T6[QR"^6_2� RF]��iTa^os"46<�]PV S&Y5T �Dta��_=T!%']Pg8g Z�V9I$e&f\S=VX[6s TQgMtQAT\]=V�]3f\lP46%7]Pg8gMZ V=��R�s�ZX]&U3[a^6s�lOVXgM4Q[QePU

�'[\Vr[\Ta]3T6e3jdZrR>]9e=YHTae3s-gMm\��Vr[QR"TQe=Uv]3e2�'R"TQgigM49��Pp ]3RFU&m

��SPR"^6_=TQV�V�46/:m �QmPK �DKaN&mM�5]9s�TgFS=R�Sve=T�eP]9S3jPT V�VrTETQg+� �6^aTU3T�R-_DZ eP]v^QT6R"]=gMm��ESPVXV T6s�ZreHUvT\g 1O^\ZrT6e=^ TQg K �DKoNDt ��� �

�3NDm

��T R"V�Zre=f=4\� mM�mn4 �QmM� m6�$[ RqR>ZXgM49[QePU�� m A2m6� T6[�APTolPm&K������&m��R"]!�)g8Zre=f$[6ePU2fvRq[a`\ZXe3f�Z eHTaV�T3�=_=[6e3s gQtos-_PT

Z�g8]9s ])�PT�R"T9^a]=R"U2]3pDYG]=UvT R-e�[Qe=U2p"]=gigMZrV��=RF]�<=]=g8^\ZrU=T6[Medgom �TQ^a]&Vr]=f=ZX[2KMJOc�t � �9L�� �3N3LDm

��Z�Tos R"Z�^6_P46/:m �5mPK��9L=JDm���S&RP�e=s&�$ZX^�APVnS=e=f=g YHTa^o_P[MedZOA$U=T g�AET�<PZXg g8T\g�U=T Rd[Qp-RFZ�Av[QePZ�gi^6_=TMe2�$[\gF_ �3R"e=T6R8m

��T6e3s-R-[ V�<PV [6sXsPp��=RD� Z ePTQR-[\VX]=f=ZXT2K��9L=J't J��3N����=c3c&m

��Z�Tos R"Z�^6_P46/:m �5mPK��9LDb=mM�$[\g _P]3R�e&R"T\gFs-T�[MSDg�U=TQY � SP[6R-s��9RP��T6S s g ^M_��q�gFs-[Qp-R>Z�A=[Qg6mM�'[\Vr[\T ]3e3s�] fvR-['�3_dZr^Q[

���'t L'bG� �3c&m

%7]=R�s"TaV�ZrSPg 4Q� m=K����vb=m6I$e=f\SDVr[6s T�^6_OTQT�As"TaTos�_'t\U=TojPTQVX]��3YGT6e3s-[ V�4\p-S=e=^6s ZX]9eP[QV�4 [aePU)Toj ]3VrS3s ZX]9eP[6R-l

Z e3s T6R"R"T\Vk[as-ZX]\ePg6m6��^6s�[ ��]3]=VX]3fPZr^Q[!%7TMe=ePZ ^a[5K � cDt K��-N �&m

A�[QS=U3R-lO4 �)m=K ���\J�� K��)�=N&ma�ePZrY [ S!)p"]=gMgiZ�VrTQg�Tosdf��6]PVr]=f3Z�T�UvT�V�W � s�s�Z�� SOT9m31=[Mj=lP46�7[QR>ZXgM4 LPN3J>�!�Dm

ATQR-[Q[QU=gM46�HmPK�� ���&m\1=SPR'V�Tag R>TQVX[os�Z ]3eOg@�=_3lOV �os�Z �\S=T\g�U=T@"$�&%� ������+����/���(6B ���� 8�� ���!(��R"[6Y$<P]3S=R-fP4&K��&b �=4

�_PZre=]=^ �QR ]=g�U\S2<$[QV�V �agMZrT6eHU=W ��Vkf��aRFZXT3m � ]3Y>� s�T\g�R"T6e=U3SOg�UvT�V�W �^a[\U��6YGZXT�UvTQg!1P^\ZrT6e=^aT\g �3c3J't ���=b��

� �=N&m

ATQR-[Q[QU=gM46�HmPK�� �vN&mM��[�p-[aS&eOT�U=Tag�U������9s g �DV �9Z g s�]=^ �aePT6g�U=T�VXW ��SDTag s U9S5V [a^ �$[as�R>]3eH�-��[6e=` [MeDZrTQ# �

Z e3s T6RM�=R��6s�[6s"Z ]3eH<PZr]=g s R-[6s"ZrfvRq[��=_PZ��9SPT3m\1P^QZXT6e=^ T6g�f��Q]=VX]3fOZ �\SPTQgM4���SPV�VXTos�Z e$LPcDt K��3N�� K �QLDm

ATQR-[Q[QU=gM46�HmPK�� ���=mM �6jPZ�g ZX]9e�U3T\g� �_=Zre=]P^aT R ]9s"ZXU3[ T� � [aYHYH[\VXZ [a#�U3S���S=R>]=V�ZXToe�U=T � Z�A=TaRqYGZ�m

�!]3YH�P[6R"[ Z�g ]3e�[ojOT\^�VrT\g�p"]3R-YGT9g�jO]&ZXgMZ ePTag6ma��e=eP[QV�T6g�U=T��'[ V �Q]3e\s-]=VX]3fOZ TEN9L7tkK ����L�K\m

Page 26: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�-�

ATQR-[Q[QU=gM46�HmPK�� ���&mM<�TQR�s �3<&R �Qg�U9S$��Z ]&^��6e=TEg SG���QR"Z�TMS=R7U9S2� � T<OT9V9,$R>TQ^6_OT6Y TQV=��R�s�g ]3S=YH[E�q��SvePZ gMZ T

^6Tae3s-R-[\VXTQ#Fm\��]3YH�=[QR-[ Z gi]9edg@<PZX]=g s-R"[Ms"Zrf=R-['�3_PZ��\SPTQgom A��6]!<OZ�]3g�J3J7t N&NvN�� �\c�K\m

ATQR-[Q[QU=gM46�HmPK����3LDmM��TQg�f=Z�g8TQYHTMe3s g�U=T!� [ Y YGZrp��6R>TQg!U\S2�*Zr]=^��MePT�gFS����6R>Z TMS=R7U=T , T6YGZ9A=V�Zks�T��=TQ4v��S=R �9SPZXT=t

LDm6 �_PZreO]3^\T6R"]9s"Z U [aT3m3�ESPV�VXTos Z eHU3S2� SPg��6SvY �$[os�Z ]3e=[ VPU=W ��ZXg s ]=ZrRFT!�$[6s�S&R"TaV�V�TQ4QL �6YHTEg��6R8m 4v��4dK �Dt �DK��

� bPm

ATQR-[Q[QU=gM46�Hmn4\[QePU� 2m3� T6R-eP]9SOg g Z-m6�"eH�=R>TQgigom6 E_OZkeO]=^QT6R"]Qs"ZXU [aT\mQ�-e �Qm � � mQ �[Mlve&[ V=[6ePU % m ��m\1�<PZk_PZ��-�VX[Q]=SPZ

�qT U3g6m #F4\�![\g [�<PV [ae=^Q[!u)��[�f=R"]9s�s T�U=TQg _PZkeO]3^ �aRF]vg6mo��T g�U3]=g giZXT6RFg�U3TEV�W ��R>^6_��Q]��-VX]3fPZXgM4 Ld4\A]9SPUvTosFm

ATQR-[Q[QU=gM46�Hmn4\[QePU)A2mM,�]3S=p"]=gomPK����=c&moI;���PTaR �CZr]=^aToedT� �_PZreO]3^QT6R>]9s"Z U [aTE� � [6YHYH[QV�Z [a#'pqR"]=Y

�7T6e3s [QVX])�v_P]3g3� K 4Q��[ ^6TQUv]9edZr[\4\A�RFTQT ^QT9m6� [QVX[QTQ]\e3s ]3f=R-['�3_PZX^6[EJ�Kac'trK\bDK�� K � �Pm

ATQR-[Q[QU=gM46�Hmn46%�m\��YH[aeDZX4 �6m � ��mQ E[olOe=[QV�4Q[QePU %�m���m916<DZr_dZ9�-�VX[Q]3SdZ�m&K������=mM��[�p-[QS=eOT�U=T ��[QYHYGZkp��aRFTQg�U9S

�7V�Z�]3^��6e=T!s"T6R"YGZ eP[QV=UPW ��_dV3[\V=��S=fv_DVr[QYG4\�![\g [�<PV [MeO^ [ 46��[QR"]=^\mQ��]=YH�3s T6g� ET6eOU9Sdg�U3TEV�W ��^6[\U��6YGZ T

U3T\g!1=^\ZrT6eO^QTQgi4 1P^QZXTMeP^6T g�U=TEV [���T6R"R"T�T6sdU=TQg��7VX[Me �os TQg �3J��Dt �=N�K�� �=N��&m

ATQR-[Q[QU=gM46�Hmn4 ��mv�VXTQYGg TQfvT6UP46�Hmo TQTaU&4 �6ma/;lve=ed4Q[6ePU��Hm\��m6 �]3YH[Me'm6�"eH�=R"T\g8g6m ��_PT �7VXTQZ�g s ]3^aT6e=T�p-[6SPe=[

pqR"]3Y �g�<O]=VXTQ43VX]��T6R'� �[ g _2<$[\VXV�TolP46�s-_PZ�])�OZ [\4Q[aeDU)Z s g�Toe=jPZXR"]3e=YHToe=s�[\V3[MePU=<PZX]3^M_PR"]3e=]3VX]3fPZX^Q[QV

Z Y$�PV�Z ^ [Ms"Z ]3e=gom A��6]!<PZX]=gMm

A�Rq[alP4 �am6�mPK�� J�KvmQ��e$s�_PT!e=[6skS&R-[\V3[aRFR-[Qe=fPTaYHToe=sd]3p&j T6R�s T'<=R"]=g T�[6ePZrYH[ V�goma��]9ePUv]9e�*TQU=Zr^Q[ VO T�O]=g Zks ] R�l

KQbdt J ����� � K c&m

A�Rq[alP4 �am6�mPK�� � �PmQ� <OgMT6R-j=[Ms"ZX]9ePg�]9e2s�_PT4�=R>TQg T6R-jPTaU)gK�=TQ^QZXYHTMePg [ae=U)g�AOT6V�Tos ]9ePg�]3p �_PZre=]3^ T6RF]9s�ZrU3[ T�Zre2s�_PT

^6]PVXVXT6^6s ZX]9eH]=p s _PT��ER>Z s�ZXg _$� SPgiToS=Y [aePU� E]9l=[ V=��]&VXVXT6f=TE]3p�1PS=R"f=Ta]3ePg 4vZkeP^ VkSOU=Zre=f s�_=TEU=T6gM^aRFZ��=s ZX]9e

]3p&s�_&R"T\T�e=T��.gK�=Ta^\ZrT\gom6� R"]=^QT6T U=Zke=fOg�]3p=s-_PT��]=]=VX]3f=ZX^6[ V&1P]=^QZXTos�l�]3p&� ]3e=U=]9e5��p"]3R K � �=N3# tXK c=c����

Kac��\JDm

A�S��6R>ZkeP4v��m K������DmK"$�&%' �����( ���G�)������ 3/ ( ��(!eO]9j'mQgK�Dmn4QS=e _DZ eP]v^ �QR-]Pg�U9S2�CZX]�� �7VXZX]3^ �MePT�U3T���S=eOZ�giZ T�U\S

��]3RFU&ma��]3^MS&Y5Toe3s g�U=TQg ��[�<=]3R"[Ms"]3ZXR"T\g�U3T�A�� ]=V ]3fOZXT�U=T�Vr[ %7[ ^6SdV s �EU3TQg!1P^\ZrT6eO^6T\g�U3T!��lO]\e K��Dt K��

b\cDm

Page 27: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

A�S��6R>ZkeP4v��m K��vN��DmM��TQg R"T\gFs-T g�U=T �_PZre=]v^ �6R>]=g�U3S)f=Z�g8ToYGT6e3sPYGZX]3^ �MePT UvT���T6eOZ9�*TaV�V [ V�46� [6R>]=^3mQA��a]PVr] fOZXT

� �aU&Z s"T6R"R-[Qe�� ToeveDT �Dt K c=b�� K c��=m

A�S��6R>ZkeP4v��m K��vN��Dm ��_=[\VXZ�^a]Osk_PT6RFZXZXU3[\T�TosP �_PZre=]v^QT6R>]9s"ZrU=[aT�����[QYHY [ V�Zr[\46�7TQR"Z�g g ]vU [ ^os�l=VX[6#7U3S2�CZr]=^��6e=T�[6S

�7V � Z gFs ]3^��MeDT�UvT�V [! �Zrp�s&<$[ V V�Tol5�q�pqR"Z �3SPT�]3R"Z�Toe=s�[\VrTQ#Fm6I�eHT T6YH�PVXT�U=W �Mjd]3VrS3s"Zr]3e't V�TEg��3S=T\VrT6sXs T4�=]Pg sM�

^6R��6edZrT6eHUvTQg-"$�9%� '����(�Tos��4 ������������� '�J���! �PV�Zr]��9�PV��\ZXg s�]=^��oePTQg6m3��SPV�VXTos"Z e�UPT�Vr[1P]=^QZ��os � f �6]=VX]3fPZ �3SOT

U3T!%�R-[ eP^6T!J�Kvt J � � ��J ���=m

A�S��6R>ZkeP4v��m K����\cDmM��TQg _PZkeO]3^ �aRF]=g�� � [6YHYH[ V�Z [ 46� T6RFZXgMg8]=U3[a^6sXlPVr[Q#'U3S2�CZX]3^��6eOT s T6R"YGZkeP[ V3[QS

�7V � Z gFs ]3^��MeDTEgFS!���6R>ZXT6S=R'T6e2 S=RF]��=TE]3^Q^QZ U&TMe3s-[ VXT9m\�!]3Y>�&[aRq[ Z�g8]\e)[6jPT ^�VXTQg�TQg�� �6^QT g [ ^os�SPTQVXV�Tag6m

��]3^6S=Y5T6e3s g�UvTQg ��[3<=]=R-[6s ]PZkR>Tag�U&TEA �a]&Vr]=f=ZXTEU3T�VX[!%'[Q^aSPVks �EU=T6g!1d^6Z�ToeO^QT6g�U=T ��lO]3eHN��Dt K�� K3K �3b=m

A�S��6R>ZkeP4v��m K����3bPmM��TQg R-_dZ eP]=^��QR-]Pg�T sdV�Tag!^M_=[\VXZ ^a]3sX_ �6R>T6g�� � [6YHYH[ V�Z [ 46� T6RFZXgMgi]=U3[a^ sXl Vk[ #'U=T\g fOZ�g8TQYHToe3s g

U3TEV [!j=[\V�V��9T�UvT�V�W ��YB]�T6e2�s�_PZX])�PZXT �����Dm)�vN�� � �)ZXe �)m �!]!�!�PTae=g�[Qe=U % m\��mM��]!�$T9V V&�-T\U3gMm #>4 ��T\g

pq[aS=ePTagD�dVXZ ]��&�PV��QZ�g s ]3^ �oeOT9g�UvT�VX[�<=[ g g T!j=[ V�V��QT�U=T�VXW ��Y ]�� �s�_PZr]!�PZ�T6# m �![a_DZrTQR"g�U=T �'[ V �Q]9e3s ]=VX]3fPZrT\4

� R"[Mj=[QS!�U=T#�=[QV��a]3e3s-]=VX]3fOZ TET6g s ��[Qp-R>Z ^Q[ Z eOT9m\��TMe s�R>T �$[Ms-Z�]9e=[\VvU=T�VX[ TQ^o_PTQR ^6_PT 1O^\ZrT6e3s"Zrp"Z �3S=T\46�'[QR"Z�gMm

A�S��6R>ZkeP4v��m K����\NDmM%7]=g giZ�V E_PZre=]=^ T6R"]9s Z U3[ T�� ��[ Y YH[ V�Z [ 4 ��T R"Z�g gM]vU [a^os�lOV [a#�p-R>]3Y ��[QT6s ]=VXZ �'���Dm��3J=c����\L��$ZXe

� mM�HmQ��T6['AOTol$[QeOU �6ma� mM�$[QR"RFZXg!�"TQU=gom #"49��[QT6s�]PVXZ�4Q[!��VXZ�]3^QTMePT�giZ s TEZ e2� ]3R�s�_PT6R"e)��[Me=`Q[MedZr[3m

��V [aRFTMe=U=]\e2�'R>TQg gi43��Pp"]3R"UDm

A�S��6R>ZkeP4v��m K����9L'mM��TQg _PZkeO]3^ �aRF]=g�� � [6YHYH[ V�Z [ 46� T6RFZXgMg8]=U3[a^6sXlPVr[Q#'U3S2� � ]3f �aePT U=TEVXW ��S=f=[ae=U=[ �'���DmaJ �����

J��vc)Zne=�$m31PTMe=S3sP[Qe=U� mM� Z ^'Avp-]=R"U)�qTaUPgom #"4vATQ]=VX]3f3l$[QePU��7[ V [ TQ])<PZX]=VX]3f3l�]3p=s-_PT ��V9<PTQR�s"ZneDT� �Zrp�s

<$[QV�V�TolO4QI$f=[QePU3[�� ��[\ZrR>T9m6�'SG<DV Z�^6[6s ZX]9e�]=^a^ [ g ZX]\e=e=TQV�V T\mQ�EToe\s-R>T � e3s"T6R"e=[6s ZX]9eP[QV!�P]\S=R�Vr[!% ]3RqY�[6s"ZX]9e�T6s

V�TQg��^a_=[6e=fvT\g�A �6]PVr]3fPZ �3SOT\g 4\��RFV �6[aePgom

A�S��6R>ZkeP4v��m6JPcvc3cDm ��_=T ��Ta]3f=TMe=T R"_PZre=]=^aT6R>]=g Tag�]=p �$[ YBZ�<PZ [\m6�'[ V [ TQ]\e3s-]=VX]3fPZr[��pqR>ZX^Q[6e=[ ���Dt K=K�� �FK�� �=m

�$[QR"RFZXgM4 �QmM� m&K �3N��&m6%7]vgMg8Z�V9 �_dZ eP]3^aTQR"]Os ZXU3[ T�� ��[ Y YH[ V�Zr[\46�7TQR"Z�gig ]vU3[\^os�lOVX[6#'p"RF]3Y �[ g s= �S=U=]=VXp-49,�T6e3lO[9m

%7]=g gMZ VO<�T6R-s T�<=R-[6s TQg�]3p'�p-RFZX^Q[�L�t KaLON � K N\JDm

�$[QR"RFZXgM4 �QmM� m&K � � �&m6%'[ YHZ�Vrl �_PZreO]3^\T6R"]9s"Z U [aT��'���'m=K��=c�� K\b �)Zre �6ma�;ma�$[6R>R"Z�g��qTQU&m #F4Q,�]=]!<PZ9%']=R-[

Page 28: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

�TQg8T6[QR"^6_2� RF]��iTa^os"46<�]PV S&Y5T J'ta��_=T!%']Pg8g Z�V9I$e&f\S=VX[6s TQgMto� R"]!<P]3g ^ Z U=Ta[\46�7TQR"Z�g8g ]vU [ ^ s�lOV [�[QePU 1vSPZXU3[\T9m

��V [aRFTMe=U=]\e2�'R>TQg gi43��Pp"]3R"UDm

�$[QR"RFZXgM4 �QmM�*49[ ePU � m A2mM��Ta[�APToldmaJPc=c��&m6��]9s�_=[af=[6Y. �_PZre=]=^QTaR"]9s"ZrU=[aT��'���'m �vN�K�� ���vb$Z e2� m A2mo��TQ[�APTol

[6ePU �6m6�*ma�$[QRqR"Z�g!�"TQU3g6m #"46� ]9s-_=[af=[6Y u5��_=T!�$['�e�]3p&�$S&YH[MePZrs�l�Zre2�[\gFs-TaRqe)�p"R"Z�^a[3m � ]=VrS=YH<dZr[

I$ePZkjOTQR"gMZ s�l�'R>TQgMg 4Q��T�� �$]3RMA&m

��Z�VX`6_OT9ZrYHTQR"4Q�;mPK��9JDb=m �#����/��!%' ����!(�(3�� 8��( % '�� 1�!/6����� �'�3� %��!/��!(�eDm g S�<=g+�'ma[6Sdg��VXUv]!�[6l ��� �DmaLPN��-N��)Z e

�HmM $TQ^�A��"T6UDm #>46/.Z�g gMTMePg ^M_=[Qp s�V�ZX^Q_=T!�RqfOT�<3ePZXg giT�U=T��VXU3]��[olC� ��PT6UPZ s�Z ]\e K���K��&m6��TMS=T!%�]vV fOTQ4

��T6p�sPJ'm��]3R"e3s-R�QfvTQR"4 ��TQZ��=`\ZXf&m

��]=]=Z �8TQR"46�Gm\�2m&K �����&ma�� E_PZ e=]v^QT6R>]3g�pqR"]=Y+s _=TEV [Ms"T �CZr]P^6T6eOT�]3p %�]3R�sd��TQR-e=[aeDm ��]v]=VX]3fPZ gi^o_PT

�*TaU=T6UvT\V Zre=fOToe$L��'t NPN�� �9JDm

��]=]=Z �8TQR"46�Gm\�2m&K ����� mM�7V�TQZXg s ]3^ ToePT!![agFsD�p-R>Z�^6[ae)R"_PZre=]=gomM% ]=g8gMZ�V9<�T6R-s T<=R"[Ms"TQg�]=pD�p"RFZ�^6[)KPt N�K������=m

��]=]=Z �8TQR"46�Hmv�2m&K �3N\JDmQ����[os�T �7V�ZX]&^aT6e=T�R-_DZneP]3^QT6R"]=g�p-RF]3Y ��[ae=fvT�<=[6[ e��TQf=4\�![��PT!�'R>]9jPZ ed^6T\mQ��e=eP[ V g�]3p

s-_OT 1O]3S3s�_��p-RFZX^Q[6e$� SPgiToSPY b��Dt K\b&K � K���K\m

��]=]=Z �8TQR"46�Gm\�2m&K �3N�� ma��U=U&Z s"Z ]3e=[QV3�*Zr]=^aTMePT s�] � VXT\Zrg s�]=^aT6ePT R"_PZre=]v^QT6R>]3g8T\g�]3p��p-R Z ^Q[\m���]=]vV ]3fPZXgi^o_OT

�*TaU=T6UvT\V Zre=fOToe$L��'tkK L���� KaN��=m

��]=]=Z �8TQR"46�Gm\�2m&K �3N=b=m6��]9s T�]\eHg ]\YGT eOT'�$V l5p-]3S=e=U;�=TQR"Z�gig ]3U=[a^os�l=V3s T\T s�_Hp�R>]3Y s�_DTE��YH])A�R"]3S!�

U3T��O]=gMZ s-gM46�s�_PZX])�PZX[9ma�'RF]=^aTQTaU&Zne=fPg�]3p&s�_DT!,�]9ePZre�APVXZ �KAPT���TaU=T6RFVr[QeOU=g T�� A3[ U=T6YGZ T!j=[ae

/.TMs-TMedg ^M_P[3���=TMeD4�� N �PtXK ���G� K��3cDm

��]=]=Z �8TQR"46�Gm\�2m&K �3N��&mM �_PZreO]=^6T6R>]9s"ZXU3[aT��'���'m��3N�K����vN �$ZXe2<Hm��Qmo� [6fPVXZ�]2[6ePU�Hm�$m\1'm\��]3]�AvT��"TQU=gom #"4

�jd]3VrS3s ZX]9eH]=pD�p"R"Z�^6[Qe$� [ Y YH[ V�gMm6�$[aR�jO[QRFUI$ePZkjOTQR"gMZ sXl�7R>TQgMg 4\� [aYH<=R>ZXU fvT3m

��]=]=Z �8TQR"46�Gm\�2m 49[aeDU$��m\1Dm\�!_=S=R"^M_DT6R8m=K����vb=m6�7T6R>Z�gig ]vU9[\^os�lOV [E]3p&s-_PTE��YG])A�RF]\S��)UvT�P]=giZks g 4Q�YGT6R"Z�^6[Qe

�!]=V�VXTa^6s-ZX]\ePgM4 ���Dm������ KvKaN)ZXe �)m � ]!�G�=T6eOg [ e=U %�m ��mo�5]!�T\VXVP�-TQU=gom #>46��T\g p-[QS=eOTQg �PV ZX]��&�dV �\Zrg s ]=^ �ae=Tag

U3TEV [#<=[\g8gMT jO[\VXV �aT�U=TEVXW ��YG])���s-_PZX]��PZ Ta# m �![Q_PZXT6R>g�UvT �'[\V��Q]\e3s ]=V ]3fPZ T94 � R"[ojP[aS!5U=T��=[ V � ]9e3s ]=VX]3f=ZXT

TQg sM��[ p-RFZX^Q[QZXe=T9m ��Toe3s�R>T �$[Ms"Z�]\e=[\V3U=TEV [! ETQ^M_PTQR-^6_PT1=^\ZrTae3s"Zrp"Z �\S=T\46�'[QR"Z�gMm

Page 29: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

��]=]=Z �8TQR"46�Gm\�2m 49[aeDU �$mM�'[6sXs"T6R>g8]3eDm=K��3N9J'mM E_DZ eP]v^6TQR"]=g8TQg�p�RF]3Y s�_dT �7V�Zr]=^aToedT�]3p&ed]3R�s�_��$Tag s�T6R-e2,�TMe lv[3m

��SPV�VXTos�Z eH]=p s _PT ��S=giToS=Y ]3p � ]3Y>�=[QR-[6s Zkj T ��]3]=VX]3f\l KQLvL't K���J �Dm

�aT6R-e3j=[ V V�4 �Qm 4\[6edU �*m6%7]3R-s T\V ZrSOg6m6JPcvc9J'm �!]3YHYH]3e)YH[6YHYH[ V�g�U3RFZrjPT s�_OT�Toj ]=VkS3s�Zr]3e=[aR�l�ZXeP^6RFTa[QgiT ]=p

_3l��Og ]3Uv]3e3sXl�Z e2s _=T ��T ]3f3Tae=T9ma�$[os S=RFT�L�KaN't b����G�qbQLPc&m

��TQ[�AOTolO4 ��m31dm��$m&K ���=b=m\�V�U\S3j=[\Z=AE] R-fOT K��=bDK�� K����DK3mM<�]=VrS=YHT)K3m �![QYH<=R"ZXU3fPT I�ePZkjOTQR"gMZ s�l�'R>TQgMgom

�![QYH<=R"Z�U3fOT9m

��Z9AOZkSPgM4Q�$m\JPc3c3JDmM��TQg�f3R"[6ePU=g�� e=fvSPV � g�U3S2� Z ]����7V�Zr]&^��6eOT�U9S���^6_=[ U)� E_DZneP]3^aT R"]3s�ZXU3[\TQ4\AZXRq[Qp"p"Z�U3[ TQ4

�![QYHTQV�ZrU3[\T6#itQg lOg s �QY)[6s"Z��\SPTQ43ZrY$�=V�Z�^6[6s ZX]9eOgD�=[ V �a]=f��a]=f9R"[��=_DZ �3SOT6g�Tos��P[QV �6]vT6e3jPZXR"]9eveDTMYGToe=s�[\VrTQgMm

��]3^6s�]=R-[\V9s-_OTQgMZXg 46I$edZ j T6R>g Zks �!�7]=Zks�Z�T6R>g 4 % Rq[aeP^QTQ4&K����8�!�Dm

��Zre3eP[QT6SOg 4v��mMj ] eDm=K N b��&m\13l g s�T6YH[!e&[Ms-S=R-[ T9m6�it\R"T6f=e3SvY [QePZrYH[\VXTOm=K c � � TQUPZ s"Zr]3eDm\1=s ]=^�A\_P]=V YG4 � J=LH�!�Dm

��]=^�A!�$]v]3U=4v��ma�)mn4Q/ ma�Bma,�ZrY <dTQV�4 [ e=U �Hm ��m � ]3_&[MePg8]9e'maJ=c=cvc&m\��T6YH�P]3R-[ V9s-R"TQe=Uvg�[6eOU2YHT6s�RFZ�^

j=[QR"ZX[os�Z�]3eHZre2sk_DT�Y [MeOU&Z <DVrT\g�[MePU)U3TMe=s-Zrs Z ]\e)]=p-�8�C(�9�+� ���KB4�N���� '%��C( � �'�6�� �/G(���(\m��a]9S=R"e=[\V3]3p=�)S=YH[ae

�jd]3VrS3s ZX]9e ���Dt J����qbvb&m

��]=]3giTQ46�Hm&K��3N&b=mM� VXT\Zrg s�]P^6T6eOT �_PZreO]=^6T6R>]9s ZXU3[QT�]=p&/ mQ�SPR-]��PT��$Zrs�_�RFT6p"TaRFT6eO^QT s�]�s�_OT�R"T9^aTMe s=s �]��

_=]=R-ePT6U�gL�PTa^\ZrT\g�]3pD�p"R8ZX^ [�[aePU 1'mM�mQ�$g ZX[9m\1d^6R>Z�� s�[EATQ]=VX]3f=ZX^6[ ���Dt K��Fb �&m

�7]=YHTQV�4Q�$mDK ���=b=m\1=SPR'V [Eg s [os�ZX]3e=�=R �o_PZXg s�]3R>Z �\S=T�UPT ��TaR-ePZ p"ZrePT\4 �PR �\g�U=T!��[ag ^a[QR-[�q��Vrf��6RFZXTQ#Fm\�gigM]3^QZX[os�ZX]9e

pqR-[ e��6[\ZXg T#�=]3S=R'V�W [Mj=[ eP^QTQYHToe3sdUvT\g�g8^\ZrT6eO^6T\g 4v�!]3YH�3s TQg� �ToeOU3S=g�KML7tkK �9LDm

�7]=YHTQV�4Q�$mDK �����=mM<�Z�giZks T p-[\Z s�T �VX[Eg s-[Ms"Z�]9e;�=R �M_PZXg s�]=R"Z �9SOT�U=T���T6R"ePZ p"ZrePT�� � [\VXZ9A3[ ]3#-�&[aR'V�T f3R>]3S��=T

TP^MS=R>gMZr]3e3ePZ�g s T �Hma�gMgi]&^aZ [Ms"Z ]3e p�Rq[ae��6[QZ�g T#�=]9SPR'VXW [6j=[ eP^6T6Y5T6e3s'UvT6g�gM^QZXTMeP^6TQgM4\�!]=YH�3s Tag ETae=U9Sdg

KaN't Jdc �G�-J�KaJDm

�7]=YHTQV�4Q�$mDK � �&b=mM��TQg�R-_DZ eP]v^ �6R"]Pg �9SP[os T6R"e=[ Z R>T6gMm\�![6R-s T�f��a]&V ] fOZ �\S=T�UPT�VXW �V f �QR"Z�TQ46�7[QV �Q]9e=s�]PVr]=f=ZXTQ4

� ]9eP]3f3R"[��3_dZrT\g 49L � �G��m

E[6lve=[\VX4 �6mI� �!m 49�Hm6��TQp-T6j=R>TQ46�Gm\AT6R"[6[ UPg 49[aePU � m6�V=A�R"[Q]3SdZ�m&K������&m �!]3e3s-R"Z�<3S=s ZX]9eHU\SHgMZ s-T

�=[\V �Q]\e3s ]=V ]3fPZ �3S=T�UvT � ZrgMg [ g p-[��-� [ g ['<PVX[MeP^Q[Q4Q� [ R"]=^6# ��S=ePT!eP]\S3jPTQV�VrT�ZXe3s-TaR��=R �os [os�Zr]3eHU\S2� Z ]��

Page 30: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

�7V�Z�]3^��6e=TEU3TEV [!�*Tag Tos [9m ��]9YH�3s�T6g ToeOU3S=g�U=T�VXW ��^ [ U��QYGZrTEU=T6g!1d^QZXToeO^QTQgM431P^\ZrT6eO^6T\g�U3TEV [E��TQR-RFT

Tos U3Tag �7Vr[Qe��6s�T g �\J��Dt ��KaN�� �\J=JDm

�_P]3U=T g 4 �m��Qm 4 �am�� ��ma �[Ml=e=[ V�4Q�HmvAT6R"[6[\U=gi49[ae=U %�m���m=1�<PZr_PZ&����V [ ]3SPZ�m=K������&m6�'R>T6YGZ �QR"TQg�U3[as�TQg� �d �O]3S=R

V�W �^Q_OT6SPV��QToe�U3S2� [6R"]=^�[os�V [ae3s"Z��9SOT��"A�R"]OsXs"T�UvT9g� �_OZkeO]=^ �QR"]vgM49�![\g8['<PV [MeO^Q[6#8m � ]=YH�3s�T g R>T6e=U3SOg�UvT

V�W �^Q[ U��MYGZ�T�U3T g 1=^ Z TMeP^QTagM4\g��6RFZ T �q� 4 ��K��DtkKPKac�� � K=KvKQbPm

�Zke=fPgFs-R � YG4\� m=K��\J3L'mo�$[ g _ � R-ePTaR'U&TaR&��Z9�G�=[QR"ZX]9e�� %�[aS3eP[!��]3R"U��q��_PZke=[ g6mM�'[\Vr[\Ta]3e3s ]=V ]3fOZ [1PZrePZX^6[

� K �-L=#Mt K�� K\b)�&m

� [ ZrT�<P46� mPK��3N3LDmM�jd]=VrS3s Zr]3e �\S=[6s�TQR-e&[ Z R"T�U3SH<=[agMgiZre)UvT�V�W � �[ g _H� �Z p�s �os-_PZX]��DZrTaeHT sP�p"[QR"# mM��]=^6s�] R-[ V

s-_OTQg ZXgM46I$eDZ j T6RFgiZks �!�7[6R>ZXg <$�>4 �7[aRFZXgM4 % Rq[aeD^ TQ4����3c8���Dm

��_PToePZrSOgM4 �mPK�� b=b=m���S=R&,�TQe3e\s�ePZ�g�U3TQRDS=e3s TQR&�dVXZX]9`��aePT6e#"$�9%� ����!(���ER�s TQeDma��ePe=[QV�Toe�U=T6R&��[os S=R-_DZXgFs-]3R"Z�g ^6_OTQg

��SPg TMS=Y /.ZXT6e��3cDt JPc3J��-J=c�� m

/.T3<dT6RF46� m K��=c\LDm6I�T3<PT6R&s�T6R�s Z �QRFT! �_dZ eP]3^aTQR"]9s-ZXUvT6e2jP]\e)UPT6R&�"ePgMT V&1&[aYH]=g6m3��SPV�V TMs"Z e�U&TEVr[21O]=^QZ�� s �

�FY$���6R>Zr[\VXT�UvTQg �$[as�S=R-[ V Z�g s�T\g�U=T � ]3gi^6]3S@K��3c3L't LPN3N��qb\c�K\m

��T6S3eOTQR&%�m&K����3LDmM��Z�T�� T6`\ZXTo_vS=e=fvT6e)`'�$Z�g ^a_=T6eB1O^M_ � U3T\Vrp"]3RqY:S&eOU ��T�<=TMeDgK�T\Zrg T#<=T\Z3U=Tae)R"T6` Toe=s�T6e)Sve=U

p"]=g gMZXV�Tae2�$[QgF_ �3R-ePT6R-eDm���T6RFZ ^6_3s"T�U3TQR&�$[Ms-S=R>p-]=R"gM^o_OT6e=U=T6e5AT6giTQV�VXg ^M_P[6p�sP`QS$%�R"T\Z�<=S=R-f �3L't J�K��"N��Dm

1PS!<=YGZ sXs"TQUJ�� � SdV l$JPc3c�� � [Q^\^aT��3s"TaU2cvc � ]9e=sX_$JPc3c9L'm

Page 31: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

� �

�!� � � ��� �H1H�!� %'�>A I ��1

%D� A I K\m��4 ��K������� �6 ��3���! 7 8�6�!�3�N���!/���%'�! �4\g�A3SPV�V3� ��� KaJ����-J&b=m ���9VX[os�T6R"[QV3jPZ�T�� �)!��3]=^Q^QVnSDg8[\V9jPZ�T�� ]3p

s-_OT�^6_OT6T'As-T Tos�_Dm91d^6[ VrT � �=c)^6Y p-]3R7�$4&K c)^QY p"]3R-�$m

%D� A I JDm6 E[os�Zr])UPZr[Qf3R"[6Y ]3p=��Z�^ T R"]3s ZrePZ&gKA9SdVXVXgom31vs-[ae=U=[6R>U � YHT6[Me5] pdV�Z jdZre&f>�4?'(3�9 ;�! +� � � J�K!s ]

J��=#Fm�"$��%� ��J�!( FC� %'���+/-��(�[QeOUH�4?)(3�� 8�! .[aR>T�T=s�[Qe3sPR>T�PR-T\g8TMe s�[6s ZkjOTQg��"A�S��6RFZreP4&K�� � cDt s-[�<dV T�J&#Fm A �$ �

KEZ�g�[#�$TQV�VN�&�&R"T g TaR�j TQUH�4?' 8���!��� ���!/���%��� .p-R>]3Y+s _PT YGZXU3UPVrT � VrT\ZXg s ]3^aT6e=TE]3p&� ] R"]=^a^Q]$���-�H1P� � �

ATQR-[Q[QU=g�[ae=U �ETQR-eP]\SPg g Z�4\Z e;�=RFTQgig #8m\1��3S=[6R>TQg�[QR"T �#?' 8������� �I��/���%��! :p-R"]=Y+s _PT ��]��TaRD�8�2[ g _'ma��_dT

j=[\VXS=TQg�]=p���?�/- ��! 8�32��3�=[6R>T p"R"] Y K c2g�A\SdVXV g�p-R"] Y.s-_PT��[\g s T6R"e2� TaU=Zks T6R"R-[QePT6[aeH� � ����74Q��8��4

��� /046���-I�C4 13���H1=#��as-_PT [ ^os�S=[\V9evS&YH<PTaR'] pdg��=Ta^\ZrYHT6ePg�p"]3R-�_DZX^6_5T6[Q^M_)YGT6[ gFS=RFT6Y5T6e3s

^Q]9SPVXU=<OT s [�APTMeHZXg fOZkj T6eHZ eH<=R"[ ^ AvT6s�g�[\p s"TaR s�_dT YGT6[ag8S3R>T6YGT6e3s=e=S=YH<OTQR"gM4 �_PZ�^o_�p"]=V�V�]!�0A S��aRFZXe

� K����vcDt s�['<PV�Tag�K�[6eOU2J �v��[3<DVrT2K # t=KE� �3#F4QJH� �3#84 �)� �\#"43LH��� #"4�� �-N3#F4��)� �3#>4&K��)�"N3#F4&K LH��L=#F4&Kvb$��LP#"4dK �

���3#>4QJ�KE�-L=#F4QJ=JH� Ka#F4QJ��)�-J=#>4 J'b$��J=#>4 �3JH��L&#Fm

%D� A I �&m\�!R"]=g g3�-giT6^6s ZX]9ePg sX_PR"]9SPf\_2s�_=TE^ R"]!�Ee)]=p=s-_PT ��JH]3pP[QeHTa[QR"Vkl �4? (��� ;�� .p"RF]3Y s�_PT�Ta[6R>Vkl

�7V�T Z g s ]=^6TMeDT�]3p=� T6ePZre��M4\� [ae3`\[MePZX[������� � AT6R"[6[\U=gM4&K���� N3# mM�[QR"V l .��$TQ[6R+0Og s�[ f=TQg�Z eHVXT6p�sd^ ]&VrS=Y)eP4

pqR"]=Y+s�]!�2s ] <O]9s�s ]3YG4\V [Ms"T6R7g s [6fPT g�Zre)R"ZXf3_=sd^6]=V SPY)eDmM�*Ta[\gFS&R"TQYHTMe3s g [aRFT!s-_OT�U=ZXg s�[6eO^6T\g@<PTosN�$TQTMe

g SP^a^QTagMg ZkjOTEg Ta^os"ZX]9ePg6m

%D� A I LDm6�'R>])�O]3R�s�Z ]3eOg�]3pPs�_dT YGTos [3�=]=U=ZX[QVXg�� � ^ �"�L� �"<�4 ��s&�"� ���-<�#�]=p'p"]=gMg Z�V�� '�+������� �6 ������! �4

^Q]3YH�=[QR"T U s�] s _=T YGTQ[Me=g�]=pDT=s�[Qe3s��#?'(3�9 �! :�"A�S��6RFZreO4dK � � c�t s-[�<dV T g�K��G��JOc&4 � K � ���3#Fma�*]=gFs

YGT s�[��=]vU&Zr[\VXg�[QR"T�<PTQVX]!�;sk_PT5K��8KEVXZre=T\4\T� ^6T��3sPYH]=g sd]=p&s-_O]3gMT�p�R>]3Y �V�U3S3jP[ Z�4'�_PZX^o_)[ R"T�g s ]9S3s T6R8m

� R>Z [Me=fPV T9g �B�[ g sP�p"R"Z�^6[��v^ Z R>^QV�Tag ��1O]3S3s�_)�p-R>Z ^Q[ �vg��3SP[6R>Tag �B��]3R-s�_)�p-RFZX^Q[\m

%D� A I b=m�"$��%' '�J�)(CBD���6 �%���E&4QYH[� Z�VXV [�� ��� KaJ���� J�K\m �^Q^QVrSPg [ V=� s ]!�=#D[QeOU)V [�<PZ [ VP�&<P]9s�s�]=Y)#DjPZ�T��g6m\1P^Q[ V�T

� Kac$^QY m

%D� A I �&m6�7]vgMZ s�ZX]9e=g��qp"R"]3Y V�T6p�s=s�]2R"Z fv_3s-#']Pp s _PT3tQ[6e3s�T6R>Z�]3R-<O] R"U=T6R ] pd]3R <PZrs 43Zre&p-R"[ ]3R <PZrs-[ V3p"]3Rq[aYGT eP4Q[QeOU

e=[\g [ V9e=]3s�^6_P4\ZXe2j=[QR"Z�]9Sdg ��ZX^QTaR"]9s"Z edZ=g�A\SPV�Vrg6m

Page 32: PLIOCENE RHINOCEROTIDAE (MAMMALIA) FROM ......$1d E ! %]=g giZ V3R"T =R>T6gMTMe9s [osZkj T6g!]3p=s-_PT!s ])T =sX[Qe3sP p"R"Z ^a[Qe)R-_dZ eP]3^QTQR]&g V Z ePTa[6fPTagM4 ! [Qe=U #"$

�9�

%D� A I N&m�"$��%' '�J�)(CBD���6 �%���E&4 �iS3j=TaePZ�VrTVrTQpXsDYH[� Z�VXV [ 4 ��� � KaJ � �XJ=cP4'�$Zks�_2���K�� � K\4\]3^ ^QVrS=g [ V9jdZ T'�)m

1P^Q[QVXT � Kac�^ Y m

%D� A I �=m ��TMS=e=T6R>W g�� K ���\L&#D[aePf=V�Tag � [ ePU�) ��Z e "$��%� '����(#F���%'�G�L/���(\m3�E]Osk_)[QR"T�ZXe2s�_=TEg [QfOZksXs-[ VC�PVX[MePT9m � �

[Qe=f=VXT#<=T6sN�TQTMe2s-_OT4�&[aRFZXT6s�[\V3[aePU2[!s�[ e=f3Tae3s=s"]_=]=R-e=<P]vg g T g �&)�� �Q[6ePf=VXT4<PT6s&��TQTMeH�=[\Vr[6s�T�[ae=U

]3^Q^ ZO�PZrs-[ V=�q]��PZ�gFsX_DZ ]3e s�] evSP^o_P[QV=^QR"TQgFs-#Fm