Estudios Geol., 55: 283·292 (1999)
THE PIG PROPOTAMOCHOERUS PALAEOCHOERUS FROM THE UPPER MIOCENE OF GRYTSIV, UKRAINE J. van der Made *, T. Krakhmalnaya ** y H. Kubiak ***
ABSTRACT
The suid from Grytsiv (= Gritsev) in Ukraine belongs to the genus Propotamochoe· rus. This genus was not yet known from the Miocene of eastem Europe. The Grytsiv suid is described and its significance is discussed within the palaeobiogeographical and stratigraphical framework of the genus Propotamochoerus. Grytsiv is geographically situated between three areas with different species of the genus: W Europe (P. palaeo· choerus), China (P. hyotherioides) and the Indian Subcontinent (P. hysudricus). In Euro· pe, P. palaeochoerus is replaced at the MN 9·10 boundary or early in MN 10 by a diffe· rent species of Propotamochoerus with affinities with P. hyotherioides or P. hysudricus. The fossils from Grytsiv extend the known range of P. palaeochoerus eastward into Ukraine. Key words: Propotamochoerus, Suidae, Mammalia, Vallesian, Miocene, Ukraine.
RESUMEN
El suido de Grytsiv (= Gritsev) en Ucrania pertenece al género Propotamochoerus. Este género todavía no se conocía en el Mioceno de Europa oriental. Se describe el suido de Grytsiv y se discute su significado dentro del marco paleobiogeográfico y estratigráfi· co del género Propotamochoerus. Grytsiv está situado entre tres áreas con diferentes especies del género: Europa occidental (P. palaeochoerus), China (P. hyotherioides), y el Subcontinente Indio (P. hysudricus). En Europa, P. palaeochoerus fue substituido en la transición MN9·1O o a MNlO por una especie distinta de Propotamochoerus con afi· nidades con P. hyotherioides o P. hysudricus. Los fósiles de Grytsiv extienden su distri· bución geográfica conocida de P. palaeochoerus por el este hasta Ucrania. Palabras clave: Propotamochoerus, Suidae, Mammalia, Vallesian, Miocene, Ukraine.
Introduction vertebrate bones. These vertebrates include small and large mammals, abundant amphibians and rep Though the loeality beeame first known in the tiles, as well as searee fishes and birds. The fo11o English literature as Gritsev (transeription from the wing mammals have been identified: Chiroptera russian), the name Grytsiv (from the ukrainian) will indet., Schizogalerix sp., Lanthanotherium sp., be used here. It is situated in the Shepetovsky dis Amphechinus sp., Domninoides sp., Proscapanus triet of the Khmelnitsky Region in Ukraine. The sp., Urotrichini spp. 1 & 2, Plesiodymilus sp., Dino fossils were diseovered in the eastern wa11 of an sorex grycivensis, Anourosoricodon sp., ?Asoricu abandoned quarry, near the village of Grytsiv in fis lus sp., Amphilagus sarmaticus, Miopetaurista sp., sure fillings in an 8 to 12 metres thiek limestone Blackia sp., Forsythia sp., Sciurotamias sp., Mono formed by a Middle Sarmatian reef. The fillings saulax sp., Palaeomys sp., Steneofiber (?) sp., Lep eonstist of elays and eontain mo11use shells and todontomys sp., Keramidomys sp., Glis vallesiensis,
* Museo Nacional de Ciencias Naturales. José Gutiérrez Abascal, 2. 28006 Madrid, España. ** Palaeontological Museum. National Museum of Natural History. National Academy of Sciences of Ukraine. 15 Bogdan Khmelnitsky street. 252601. Kiev 30, Ukraine. *** Instytut Systematyki i Ewolucji Zwierat Polskiej Akademii Nauk, u!. Slawkowska 17.31016 Kraków, Poland. 284 J. VAN DER MADE, T. KRAKHMALNAYA, H. KUBIAK
Muscardinus topachevskii, Myoglis ucranicus, 20 * Paraglirulus cf. werenfelsi, Miodyromys gryciven 18 Cm sis, Anomalomys sp., Lophocricetus sp., Cricetulo * 16 don complicidens, Sarmatomys podolicus, Allohye * na sarmatica, Eomellivora wimani piveteaui, Ictit .5 14 x * herium spelaeum, Mustelidae indet., Gomphothe 12 rium sp., Rhinocerotidae indet., Chalicotherium sp., ... Hipparion primigenium, Hipparion sp., Dorcathe 10 rium sp., Lagomericinae sp., Dystychoceras sp., 8 8 10 12 14 16 18 Euprox sp., Procervulus sp., Cervulinae sp. (Korot Po kevich, 1998; Korotkevich et al., 1985; Nesin & Fig. l.-Bivariate plot of the male lower canine. Legend and Kowalski, 1997; Rzebic-Kowalska & Topachevsky, provenance of data as in Figure 2. 1997; Semenov, 1989, 1994; Wolsan & Semenov, 1996). The presence of Hipparion and the absence of murids, suggests an MN9 age. IPUW Institut für Paliiontologie der Universitiit, Wien. Apart from sorne short descriptions of the Hippa IVAU Instituut voor Aardwetenschappen, Utrecht. rion (Krakhmalnaya, 1994, 1996a, 1996b), none of IVPP Institute for Vertebrate Paleontology and Paleo the large mammals from Grytsiv have been descri anthopology, Academia Sinica, Beijing. MAMA Museu Arqueologic Municipal "Camil Visedo bed. Korotkevich (1988) identified sorne suid Moltó», Alcoy. remains from Grytsiv as ?Hyotherium sp. Besides, MGL Museum Guimet, Lyon. Microstonyx has been described from the Miocene MNCN Museo Nacional de Ciencias Naturales, Madrid. MNHNP Muséum National d'Histoire Naturelle, Paris. and Propotamochoerus from the Pliocene of Ukrai NMB Naturhistorisches Museum, Base!. ne (Korotkevich, 1967, 1970, 1976, 1988; Krakh NMM Naturhistorisches Museum, Mainz. malnaya, 1989). The suid material from Grytsiv has NMW Naturhistorisches Museum, Wien. MPV Museo Paleontológico de Valencia. increased and it is the aim of this paper to describe PIMUZ PaH¡ontologisches Institut und Museum der Uni it, assign it to a taxon and discuss its palaeogeo versitiit, Zürich. graphical significance. SLJG Steiermiirkisches Landesmuseum Joanneum, Graz.
Measurements and terminology Systematics All measurements in mm and taken as indicated by Van der Made (1996). Order Artiodactyla Owen, 1841 Nomenc1ature of the tooth morphology from Van der Made (1996). Superfamily Suoidea Gray, 1821 DAP Anteroposterior diametre of a tooth or bone. Family Suidae Gray, 1821 DAPd DAP at the distal end of abone. Subfamily Suinae Gray, 1821 DAPp DAP of the proximal end of abone. Genus Propotamochoerus Pilgrim, 1925 DLL Labio-lingual diametre of an incisor. DMD Mesio-distal diametre of an incisor. Species Propotamochoerus palaeochoerus DT Transverse diametre. (Kaup, 1833) DTa DT of the anterior lobe of a tooth. DTp DT of the posterior lobe of a tooth or proximal end of abone. DTpp DT of the third lobe of a M3. Selected synonymy DTd DT of the distal end of abone. DTdf DT of the articular facet at the distal end of abone. L Length of abone. 1833 Sus palaeochoerus Kaup - Kaup: 11-13, pI. 1, figs. 1-3. 1926 Sus (Hyotherium) palaeochoerus - Pilgrim: Collections and their abbreviations 11, pI. 1. 1971 Korynochoerus palaeochoerus (Kaup) The fossils are housed in the Department of Vertebrate Pala 1833 - Schmidt-Kittler: 129-168. eozoology and Paleontological Museum of the National 1992 P. [ropotamochoerus] palaeochoerus - Van Museum of Natural History, Kiev (NMNHK) and were compa red with fossils that are kept in the following institutions. der Made, Montoya & Alcalá: 402, 406, BSPHGM Bayerische Staatssammlung für PaH¡ontologie 411. und historische Geologie, München. Type locality: Eppelsheim. GSP Geological Survey of Pakistan, Islamabad. HLD Hessisches Landesmuseum, Darmstadt. Age of type locality: Early Late Miocene, Valle 1M Indian Museum, Calcutta. sian, MN9. IPS Instituto de Paleontología, Sabadell. Material: All material is indicated in tables 1-3. THE PIG PROPOTAMOCHOERUS PALAEOCHOERUS 285
38 ... Grytsiv 18 20 M x 4 ... 36 3 p 18 MI '1- \ P. palaeochoerus 16 ..t:x o"~o * x*~' +++ Q. + MNll·13 !i 14 !i 16 +"'" .t: 34 1 i",:*x o o ,. « pX' Xtt' X P. hysudricus 0 32 ,~o'\f:o ' 12 x~:X 14 '!ft~ * P. hyolherioides X 10 12 30 -¡. 10 12 14 16 18 20 10 12 14 16 18 20 22 28 DT DTa 12 14 16 18 20 22 DTa 30 38 2 3 28 22 28 M 36 M * M P ~ 26 2 20 4 26 34 * * * .; -: + Q. 24 Q. 18 - 24 * 32 '+ « « *~~* ,,~";, *+ 0 Jj~ 0 22 16 ~o !i 22 ° 0 .+ !i 30 ~: ~ .. o O O )4" + ~ :,o.¡., x~o+ X + + 20 XX 14 ~ 20 Xx.- ' 28 • X.o • x X +""l<. 18 12 18 26 10 12 14 16 18 20 6 8 10 12 14 16 DTp DTp 16 24
24 22 14 22 14 16 18 20 22 24 16 18 20 22 24 26 28 P 22 MI 20 3 DTa DTa Fig. 3.-Bivariate plots of the upper cheek teeth. Legend and Q. 20 01 Q. 18 x. ,+, « 01 •t< « provenance of data as in Figure 2. 0 18 .,:1".: 0 16 9-:: X *t,W )t:, * 16 + * 14 + 20,------, 24,------, 20,------, 14 12+---,-....~.----,---1 8 10 12 14 16 18 4 6 8 10 12 14 18 22 18 DTp DTp * 20 16 i¡. • Fig. 2.-Bivariate plots of the lower cheek teeth. Propotamo O ...." §l18 :::;; 14 + choerus palaeochoerus from Grytsiv (NMNHK), P. palaeocho + o o + erus from various localities in Europe (BSPHGM, HLD, IPS, ... IPUW, MGL, MNHNP, NMM, NMW, SLJG), Propotamocho 12 16 12 erus sp. of MN 11-13 from Europe (MNCN, MPV, NMB, 10 14 10 NMW), P. hysudricus from various localities in Pakistan (GSP, 1M, IVAU), P. hyotherioides from Lufeng (IVPP; Van der 8+-,.....,..~,...... ,.-I 12+T'"'""l~""""""'¡ 8+...... ~..,.....,--1 Made & Han, 1994). 6 8 10 12 4 6 8 10 2 4 6 8 DLL DLL DLL Fig. 4.-Bivariate plots of the upper incisors. Legend and pro venance of data as in Figure 2. Description and comparison
The 11 (PI. 1, fig. 7) and 12 (PI. 1, fig. 8) have crowns of an to be narrow. P. palaeochoerus (dots in fig. 1) and the intermediate height. In Hyotheriinae and Cainochoerinae, the MN 11-13 Propotamochoerus (square) tend to have both crowns are lower, in many Tetraconodontinae and Suinae, the sides equally wide or even the posterior side wider, whereas crowns are of similar height, but in part of the Suinae, inclu P. hysudricus (triangle) and P. hyotheriodes (stars) tend to ding Sus the crowns are higher. In the incisors with higher have wider labial sides « Table I.-Measurements (in mm) of the associated cheek teeth ofPropotamochoerus paÚleochoerus from Grytsiv P2 P3 P4 MI M2 M3 22-1067 R DAP 14.4 15.8 16.5 18.2 21.9 31.2 DTa 5.3 7.1 9.8 12.4 14.9 16.8 DTp 6.2 8.4 11.6 12.9 15.2 14.7 DTpp 11.6 22-1068 L DAP 14.0 16.4 16.6 18.1 22.0 31.2 DTa 5.2 7.1 9.8 12.3 14.8 16.5 DTp 6.0 8.3 11.7 12.7 15.0 14.8 DTpp 11.6 22-1071 (M3) R DAP 22.5 29.3 22-1072 DTa 15.7 16.2 DTp 15.7 14.4 DTpp 11.7 p2 p 3 p4 MI M2 M3 22-1064 R DAP 14.2 18.6 22.4 29.3 DTa 16.5 16.8 19.1 19.6 DTp 15.8 18.1 17.5 DTpp 10.2 22-1063 L DAP 14.9 18.4 22.6 29.2 22-1063a (M3) DTa 16.7 16.7 19.9 19.7 DTp 15.7 18.2 16.9 DTpp 11.1 known in these teeth. The two islolated teeth are tentatively cusp, there is also a pentapreconule and sorne lateral wrinkles assigned to P. palaeochoerus, but the possibility that they (formed by the pentaectocrista and pentaendocrista). belong to one of the small Parachleuastochoerus species Sorne deciduous teeth and bones have the common suid cannot be excluded. morphology. The P2 and P3 (PI. 2, figs. 1 & 4) have a flattened protoconid In figures 2 & 3 the DAP and DT of the teeth of the different with anterior and posterior crests. In the Pf in the mandibles, species of Propotamochoerus. P. palaeochoerus (dots) and the the protopostcristid is a better developed than in the isolated Grytsiv (triangles) suid tend to have large first molars and P4, premolars. The P4 (PI. 2, figs. 1 & 4) has a metaconid that is while the second molars are not so big, and the third molars are placed close to and slightly behind the protoconid. Though the even smalI compared to the other species. The relative size of differences are small, the cusp is better developed than in the the third (and to sorne extend the second) molar reflects diet; majority of the species of this genus and is similar in size and larger or longer M3 suggest a more herbivorous diet. P. hyothe position to the metaconid of the P. palaeochoerus P4• The pro rioides (stars), P. hysudricus (oblique crosses) and the Euro toprecristid becomes rapidly lower away from the protoconid. pean Propotamochoerus of MN 11-13 (crosses) seem to have The talonid has a well developed hypoconid, more or Iess in the relatively large M3 and smalI P4 compared to P. palaeochoerus middle. The tooth is of the Dicoryphochoerine type (sensu (dots). Schmidt-Kittler, 1971 and Van der Made & Moya-Sola, 1989). The MI and M2 (PI. 2, figs. 1 & 4) have the common suine morphology. The M3 (PI. 1, fig. 9; PI. 2, figs. 1 & 4) has a third lobe consisting of a pentaconid in the middle, preceeded by a Discussion pentapreconulid and several smalI cusplets and wrinkles of the enamel to the sides (formed by the pentaendocristid and penta ectocristid). A specimen from Ravin des Zouaves has a penta The Vallesian suids inelude very small Cainochoer conid and hexaconid (De Bonis & Bouvrain, 1996, fig. 9), whe inae, lophodont Listriodontinae, Tetraconodontinae ras other MN 11-13 Propotamochoerus do not have a hexaco with an enlarged P4 with only one main cusp, nido Hexaconids are known in the M3 of P. hyotherioides, but not in those of P. palaeochoerus and P. hysudricus. Chinese Hyotheriinae with no paraendoconulid in The Cr (PI. 1, fig. 4) has a 10w crown and a rapidly narro the p4 and Suinae of the tribe Dicoryphochoerini. wing rool. The morphology of the premolars suggests that the The p4 (PI. 1, fig.l; PI. 2, fig. 2) has the paracone and meta cone welI separated. The paraendoconulid and metaendoconu Grytsiv suid belongs to the Dicoryphochoerini, a lid are clearly present, though not big. tribe of the Suinae (Schmidt-Kittler, 1971; Van der The MI and M2 (PI. 1, fig. 1) have the typical suine morpho Made & Moya-Sola, 1989). In a recent elassifica logy, though the crests or lobes are not as welI visible as in tion, this tribe ineludes: Propotamochoerus (=Kory several of the younger genera of the Suinae. The M3 (PI. 1, fig. 2) has a simple third lobe consisting of a pentacone that is nochoerus), Hippopotamodon (=Microstonyx), placed lingualIy of the axis of the tooth. In addition to this Eumaiochoerus, Kolpochoerus and Hylochoerus THE PIG PROPOTAMOCHOERUS PALAEOCHOERUS 287 Table 2.-Measurements of the lsolated teeth of intially been assigned to Sus and later to Hyothe Propotamochoerus palaeochoerus from Grytslv rium and Korynochoerus. Pickford (1988) stated that the species was very similar to Propotamochoer DMD DLL us hysudricus and Van der Made et al. (1992) were 22-1079 1, L 6.6 9.9 possibly the first to place the specíes formally in 22-1077 1, R 6.7 10.0 Propotamochoerus, though the justification for that 22-1078 11 R 6.6 10.7 remained a long time in press (Fortelius et. al., 22-1080 12 R 8.5 11.5 1996). The genus Propotamochoerus includes the 22-167 12 R 8.4 12.5 22-1081 1 L 6.5 10.2 following species: P. palaeochoerus, P. hysudricus, 3 P. hyotherioides, P. wui, P. provincialis and a form 22-1053 13 R 6.5 9.6 22-1082 13 R 6.5 9.4 that entered Europe either in MN 10 or MN 11 and 22-379 13 R 6.3 10.5 that lived on till at least MN 13 (Pickford, 1988; 22-2914 Dl2 L 4.9 8.3 Van der Made & Moya-Sola, 1989; Van der Made & Han, 1994; Fortelius et al., 1996). The limits of DAP DTa DTp DTpp the geographical distribution of these species is not 22-951 Cf L 12.4 7.4 known. Pliocene P. provincialis has been described 22-2917 D3 R 14.9 7.0 11.4 from Kuchurgan (Korotkevich, 1967, 1988) and 22-2918 D4 L > 12.7 Kvabevi (Vekua, 1972). Miocene Propotamochoer 22-1025 p4 R 14.0 16.0 us have not been cited from the area and the 22-1073 M' L 18.9 17.1 15.7 collection from Grytsiv is the first one in being des 22-1074 M' R 19.0 16.9 15.6 22-1070 M2 R 22.5 19.8 18.7 cribed from the area that extends from the north of 22-158 M3 L 29.7 20.9 19.0 12.0 the Black Sea to the north of China. Grytsiv is thus 22-2916 P, R 9.6 4.0 4.2 in a position between the three areas from where the 22-10.5 P2 R 12.8 4.7 5.4 genus is well known. 22-1076 P4 R 10.9 P. wui is a very small species from the Late Mio 22-1058 M2 R cene of China (Van der Made & Han, 1994) and is Li La Po 22-2915 Cm L 11.0 10.5 10.1 much smaller than the Grytsiv suid. P. hysudricus. Pickford (1988) revised the Mio cene Suids of the Indian Subcontinent and recog nized only P. hysudricus. Skulls assigned to this (Van der Made, 1997). All specíes of Hippopotamo specíes have either the parietal ridges wide apart don are much larger than the Grytsiv suid. Eumaio and a straight dorsal profile or have these crests choerus is an endemic of a Miocene island that close together and a concave profile with the occi included Toscane and Sardinia and has a number of pitals appearing as being elevated. It is the ques peculiar features. Kolpochoerus and Hylochoerus tion, whether all this material represents one spe are known only from Africa and are suids with a cíes, or whether there is a rupture like in Europe. highly specialized dentition, with incisors with Until this problem is resolved, we follow Pickford increased DMD relative to DLL, with very elonga (1988) and treat the material as representing one ted M3 and with sorne tendency to increase hypso specíes. donty. The suid from Grytsiv belongs to Propota The material assigned here to P. hysudricus inclu mochoerus. des tooth rows with premolars, sorne isolated pre Most of the species of Propotamochoerus were molars, but no isolated molars. The material is pre originally assigned to the genus Sus. This is for in dominantly from Dhok Pathan equivalent strata, or stance the case with P. palaeochoerus, which has of unknown exact provenance. M3 tend to have a Table 3.-Measurements of the bones ofPropotamochoerus paloeochoerus from Grytslv Humeros >DAPd DTd DTdf RI R2 R3 R4 R5 22-1069 R >48.9 54.4 39.5 33.6 24.9 26.3 24.4 25.2 MP I1/MP v DAPd DTd 22-119 L 14.8 9.3 22-1087 L 16.4 11.9 Lateral phalanx 3 DApp DAppo DTp L 22-395 R 9.7 8.7 7.9 > 17.9 288 J. VAN DER MADE. T. KRAKHMALNAYA. H. KUBlAK Plale I.-Propowmochouu$ pafoeoclwf'rlls fmm Grylsiv. Fig. 1.-22-1067 - righl mandible wilh P~-MJ' a) lingual view. b) ocelu sal "tew. e) buttal vicw. Fig. 2.-22-1025 - right 1'"; a) bucealllod b) occlusal views. ng. 3.-22-2916· righl PI; al occlusal. b) lingual. and e) buceal views. Fig. 4.-22-1068 - left mandiblc Wilh P~-MJ' a) occlusal and b) lingual vicws. Fig. 5.-22-1085 • righl P:; a) lingual. b) buceal. aOO e) occJusal vieW5. 1ñc bar represenl~ 2.5 cm for figs. 2. 3 & 5 and 5 cm for figs. I & 4. THE PIG PROPOTAMOCIIOERUS PALAEOCHOERUS 289 P1a!e 2.-PrQPowmochwrtu palu~och(Nrwfrom GrylSiv. Fig. 1.-22-1064 - righl maxillll wilh P'_Ml fully rrupled and wilh Ml in alveolus; a) buccal and b) occ1usal Vlc:ws. Fig. 2.-22-158 - left Ml; occlusal view. Fig. 3.-22-1068 (delail) - Ieft P~-MJ; occlusal Vlc:w. Fig. 4.-22-951 -Iert C: a) linsua!. b) bucea!. and e) occlusal views. Fig. 5.-22-2915 -Ief! C-; a) postero-infe 7.~i2-1079 rior. and b) buceal View5. Fig. 6.-22-1053 - ngbt 13: a) mes)o.-lingua!. b) labial. and e) distal vieW5. Fig. ·Ieft It; _a) dista!. b) lin¡ual. and e) mesial vicws. Fig. 8.-22-1080 - right 11; a) mesia!. and b) lingual view5. Ag. 9.-22-1071 - right Ml; a occlusal and b) lingual views. 1be bar represenlS approximately 3.6 cm for [jgs. 2-9 and 5 cm for fig. 1. 290 J. VAN DER MADE, T. KRAKHMALNAYA, H. KUBIAK simple third lobe, but is relatively large. Whereas became extinct after MN 15. P. hyotherioides, P. the MI and P4 tend to be smaller than in P. palaeo provincialis and the MN 11-13 form are morpholo choerus, the M3, and in particular the M3 tend to be gically very similar, though the latter is the most large. The P4 and MI from Grytsiv are within advanced in 12 elongation. The relation of these (lowers) or aboye (upper teeth) of the metrical ran three forms with the Indian one(s) is not clear. P. ges of P. hysudricus, while the M3 are in the lower provincialis differs from the Grytsiv form in size ranges. Not only the size differs, but also the pro and in tooth proportions. portions. P. palaeochoerus apeared in Europe in MN 8 The MN 11·13 Propotamochoerus from Europe (or late MN7+8). The genus is particularly abun was initially placed in P. palaeochoerus, but is now dant in MN 9, but became rare or extinct in recognized to be different and is believed to have MN 10. The species has a concave skull profile, affinities with P. hyotherioides (Fortelius et al., inflated zygomatic arcs, incisors with not very 1996) or P. hysudricus (De Bonis & Bouvrain, high crowns, an 11 with a large distal cusp, not 1996). It replaced P. palaeochoerus at the MN 9-10 very large or elongate 12-3, small and short cm, transition or early in MN 10. The genus is rare in small Cm with a particular section, M3 with a pen MN 10, which explains this incertainty. A juvenile taconid in the middle and no hexaconid, P4 with a skull and mandible are known from Samos relatively well developed metaconid and propor (< References Kubiak, H. (1981). Suidae and Tayassuidae (Artiodacty la, Mammalia) from the Miocene of Przeworno in Bonis, L. de & G. Bouvrain (1996). Suidae du Miocene Lower Silesia. Acta Geol. Polonica, 31: 59-70, 3 pia supérieur de Grece. Bull. Muséum Nat. d'Histoire tes. Naturelle. n.º 1, 4e série, 18, section c, Paris: 107-132. Made, J. van der (1996a). Listriodontinae (Suidae, Mam Dam, J. A. van (1997). The small mamma1s from the malia), their evolution, systematics and distribution in Upper Miocene of the Terue1-A1fambra region (Spain): time and space. Contrib. Tertiary Quaternary Geol., pa1aeobiodogy and paleoc1imatic reconstrucitons. Geo 33: 3-254, 19 tables. logica Ultraiectina, 156, 204 pp. - (1996b). Pre-Pleistocene land mammals from Crete. Forte1ius, M., Van der Made, J. & Bernor, R. L. In: Pleistocene and Holocene Fauna 01 Crete and its (1996). Midd1e and Late Miocene Suoidea of Cen First Settlers (D. S. Reese, ed.) Monographs in World tral Europe and the Eastern Mediterranean: Evo1u Achaeology, Madison, 28: 69-79. tion, Biogeography, and Pa1eoeco1ogy. In: The Evo - (1997a). On Bunolistriodon (= Eurolistriodon) and lution 01 Western Eurasian Neogene Mammal Fau kubanochoeres. Proceedings 01 the Koninklijke nas. (R. L. Bernor, V. Fah1busch & H. W. Mitt Nederlandse Akademie van Wetenschappen, 100: mann, eds.), Columbia University Press, New York: 141-160. 348-377. - (l997b). Intercontinental dispersa1 events, eustatic Haq, B. U., Hardenbo1, J. & Vai1, P. R. (1987). Chrono sea level and Early and Middle Miocene Strati logy of F1uctuating Sea Leve1s Since the Triassic. graphy. In: Actes du Congres BiochroM'97 (J. P. Science, 235: 1156-1166. Agui1ar, S. Legendre & J. Micheax, eds.), Mém. et Hellmund, M. (1995). The vertebrate locality Maramena Traveaux EPHE de [,Institut de Montpellier, 21: (Macedonia, Greece) at the Turo1ian-Ruscinian Boun 75-81. dary (Neogene) 13. Suidae (Artiodacty1a, Mamma1ia). Made,1. van der & Han Defen (1994). The Suoidea from Münchener Geowissenschaltliche Abhandlungen, A, the hominoid locality Lufeng (Yunnan, China). Proce 28: 143-156. edings 01 the Konink[¡jke Nederlandse Akademie van Kaup, J. J. (1833). Description d' ossements lossiles de Wetenschappen, 97: 27-82. Mammlferes 2. J. G. Heyer, Darmstadt: 1-31. Made, J. van der Montoya, P. & Alcalá, L. (1992). Korotkevich, E. L. (1967). Fauna of the 1arge mamma1s Microstonyx (Suidae, Mammalia) from the upper Mio from the P1iocene deposists of the Kuchurgan va1ey. cene of Spain. Geobios, 25: 395-413. In: Mesto i znachenie iskopaemych mlekopitajuzshix Made, 1. van der & Moya-Sola, S. (1989). European Sui Moldavii v kainozoe SSSR. Kishinev: 77-85 (in Rus nae (Artiodactyla) from the Late Miocene onwards. sian). Boll. Soco Paleontol. Italiana, 28: 329-339. - (1970). Mamma1s of Beris1av old Sarmatian fauna. Nesin, V. & Kowa1ski, K. (1997). Miocene Gliridae In: Priodnaya obstanovka i launy proshlogo, Naukova (Mamma1ia: Rodentia) from Grytsiv (Ukraine). Acta Dumka, Kiev, 5: 24-121 (in Russian). Zool. Cracoviense, 40: 209-222. - (1976). The main localities of Hipparion fauna on the Pevzner, M. A. & Vangengeim, E. A. (1993). Magne territory of the Northern Black Sea coast area. Vestnik tochrono1ogical age assignements of Middle and Late Zoologii, 6: 65-72 (in Russian). Sarmatian Mammalian localities of the Eastern Para - (1988). A history of the Hipparion fauna of Eastern tethys. Newsletters on Stratigraphy, 29: 63-75. Europe. Kiev, Naukova Dumka, 106 pp. (in Russian). Pickford, M. (1988). Revison of the Miocene Suidae of Korotkevich, E. L., Kushniruk, V. N., Yu. A. Semenov t~e Indian Subcontinent. Münchener Geowissenschalt & Chepa1yga, A. L. (1985). New locality of the Middle [¡che Abhandlungen, Reihe A, Geologie und Paliionto Sarmatian vertebrates in the Ukraine. Vestnik zoologii, logie, 12: 1-91. 3: 81-82. Pilgrim, G. E. (1926). The fossil Suidae of India. Krakhma1naya, T. V. (1989). Microstonyx major (Artio Memoirs 01 the Geological Survey 01 India, n. ser., 8: dacyla, Suidae) of early Meotian Fauna of Novaya 1-65. Emetovka. Deponirovano v VINITI, N. 4289-v 89, Rzebic-Kowalska, B. & Topachevsky, V. (1997). Insecti 20 pp. (in Russian). vora (Mammalia) from the Miocene of Grytsiv in - (1994). Hipparions from the old Sarmatian fauna of Ukraine. I. Heterosoricidae Viret & Zapfe (1951. Acta ~he Northern B1ack Sea coast area. In: Stratigraph Zool. Cracoviense, 40: 237-247). lchny ta paleontologichny doslidzhennaya v Ukraini. Schmidt-Kittler, N. (1971). Die obermioziine Fossilla Kiev, Institut geologischeskihk nayk Press: 64-66 (in gerstiitte Sandelzhausen 3. Suidae, Artiodactyla, Mam Russian). malia. Mitteilungen der Bayerischen Staatssammlung - (l996a). Hipparions of the Northern Black Sea coast lür Paliiontologie und historische Geologie, 11: 129 area (Ukraine and Moldova): species composition and 170. stratigraphic distribution. Acta zool. Cracoviense, 39: Semenov, Yu. A. (1989). Ictitheres and morpholically 261-267. similar hyaenas from the Neogene of the USSR. Nau - (1996b). Hipparion fauna of the early Meotian of the kova Dumka, Kiev: 180 pp. (in Russian, English abs northern B1ack Sea coast area. Naukova Dumka, Kiev, tract). 226 pp. (in Russian, with Eng1ish abstract). - (1994). Allohyaena sarmatica (Carnivora, Mammala) Krakhma1naya, T. V., Svetlitskaya, T. V. & Chepalyga, - a new hyaenid species from the late Miocene of the A. L. (1993). New data on stratigraphy, magnetostrati Ukraine. Acta zoologica cracoviense, 37: 31-38. graphy and mammal faunas of the late Miocene loca T~enius, E. (1950). Postpotamochoerus n. subg. hyother litY of Novaya Emetovka (Ukrania). Newsletters on IOdes aus dem Unterplioziinvon Samos (Griechenland) Stratigraphy, 29: 77-89. und die Herkunft der Potamochoeren. Sitzungsberichte 292 J. VAN DER MADE, T. KRAKHMALNAYA, H. KUBIAK der Osterreichische Akademie von Wissenschaften, mat Middle Sarmatian micortheriofauna (lnsectivora, hematisch-naturwissenschaftliche Klase 1, 159: 25-36. Lagomorpha, Rodentia) in the Eastern Europe. Dopo Topachevsky, V. A., Nesin, V. A. & Topachevsky, 1. V. vidi natsional'noj Academii Nauk Ukrainy, Paleonto (1997). The essay of the microtheriofauna history logy, 2: 107-110 (in Russian). (lnsectivora, Lagomorpha, Rodentia) of Ukraine Vekua, A. K. (1972). Kvabebi fauna of the akchagyl ver during Middle Sarmatian-Akchagyl periodo Vestnik tebrates. Nauka, 351 pp. & 36 plates, Moscow (in Rus zoologii, 31: 3-14 (in Russian). sian with English abstract). - (1998). Biozonal microtheriological scheme (stratigraphic Wolsan, M. & Semenov, YU. A. (1996). A revision of distribution of small marnmals - Insectivora, Lagompha, the late Miocene mustelid carnivoran Eomellivora. Rodentia) of the Neogene of the northem part of the Eas Acta 2001. Cracoviense, 39: 593-604. tem Paratethys. Vestnik zoologü, 32: 76-87 (in Russian). Topachevsky, V. A., Nesin, V. A., Topachevsky, 1. V. & Recihido el15 de junio de 1999. Semenov, Yu. A. (1996). The oldest locality of the Aceptado el19 de octubre de 1999.