A review of Bulgarian Hipparion: Mammalia, Perissodactyla

A REVIEW OF BULGARIAN HIPPARION (MAMMALIA , PERISSODA CTYLA) by ANN FORSTEN * RI~SHMI~ ABSTRACT Les esp~ces du genre fossile Hipparion trouv4es jusqu'h pr4sent en Bulgarie sont pass4es en revue ainsi que leurs affinit4s. Parmi les 8 esp4ces diff4- rentes qui avaient 4t4 d4crites, 3 seulement sont consid4r4es comme valides. The species of the fossil genus Hipparion found in Bulgaria to this date, and their affinities, are discussed. Of the earlier described 8 different species, only 3 are here considered possibly valid. MOTS-CLt~S : SYNTHI~SE BIBLIOGRAPHIQUE, HIPPOMORPHA, NI~.OGI~NE, GITE FOSSILIFI~RE, NOMENCLATURE, BULGARIE. KEY WORDS : REVIEW ARTICLE, HIPPOMORPHA, NEOGENE, FOSSILIFEROHS DEPOSIT, NOMENCLATURE, BULGARIA. * Zoological Institute, University of Helsinki, Helsinki, Finland. G4obios, n ° 11, fasc. 1 p. 31-41, 5 fig., 12 tabl. Lyon, f4vrier 1978 --- 32 - - INTRODUCTION I. Nikolov in an unpublished monograph recently described the Bulgarian hipparions. Some of the main points in this paper have been published (1971, 1973a, 1973b). Hipparions were earlier more briefly discussed in the large study on Bulga- rian mammalian fossils by P. Bakalov ~ I. Nikolov (1962). The material described by I. Nikolov (1973b) derives from some 30 fossils localities in different parts of Bulgaria, believed to span a stratigraphic range from the Middle Sarmatian to and including the Post-Pontian Levantine. The most important sites as to their amont of fossils are Nesebr, Kalimansti, and Kromidovo (fig. 1). The fauna of Nesebr derives from the Middle Sarmatian, whereas the two latter derives from the Meotian to Middle Pontian inclusive. These Hipparion,Lfaunas must be counted among the im- portant ones in Eurasia. Geographical,ly they connect the classical faunas of SW U.S.S.R. and those of Central and South Europe. I. Nikolov (1973b) distinguishes a total of 8 Hipparion species, of which 3 new, in addition to two unnamed forms. A review of the Bulgarian material indicates that considerably fewer taxa are, in fact, represented. My thanks are due to the Embassy of the Bulga- rian Socialist People's Republic in Helsin:ki for helping to arrange my visit to Sofia in 1976, to Dr. I. Nikolov for handing over to me the inte- Abbreviations used in the diagrams: E = Ezerovo, H Hadjudimovo, I=Ivanane, K=Kalimantsi, Kr=Kromidovo. resting material of fossil horses which he personally excavated, curated, and worked, and to the staff at the Paleontological Museum of the University of Sofia for their help and many kindnesses. The h~pparion material, which consists of a few skulls, several jaws, tooth rows, isolated teeth, and limb bones, is housed in the Paleontological Museum. aa 6 Fig. 1 - Carte de la Bulgarie montrant la situation des gisements cit6s dans ie texte. Map of the Bulgarian People's Socialist Republic showing fossil sites referred to in the text. (From Atlas Mira, 1967). 1 : Nesebr 4 : Ploski Blagoevradsko 7 : Ivanane 2 : Kalimantsi 5 : Ezerovo • 8 : Hrabrsko 3 : Kromidovo 6 : Hadjidimovo DESCRIPTION Nesebr In the fauna from the Middle Sarmatian of Nesebr on the Black Sea coast I. Nikolov (1973b) identifies H. nesebricum BAKALOV ~ NIKOLOV, H. mediterraneum ROTH & WAGNER, and H. pra,esul, catum NIKOLOV. The distinguishing characteristics of these species, as delimited by Ni,kolov, are not quite clear. All are said to be rather large and massive forms the teeth of H. nesebricum and H. mediterraneum little to moderately plicated, those of H. praesultacum strongly plicated. The proto- cone of the teeth of H. nesebricum is described as elongated and flattened, and I. Nikolov (p. 9) compares the teeth in this respect with those of some N. American hipparions. ' The protocone of .58 .54 .50~ .46 ! ~r KL K / Kr MC III I/dist.br - - 33 - .62 .58 .54 .50 . .46. r .~8 .~2 .~6 .3'6 .4o' t Fig. 2 - Diagramme de dispersion de la largeur distale articulaire des MC III en fonction de la longueur totale (6ch. logarithmique). Les ellipses d'6quiprobabilit6 ~ 95 % ont 6t6 dessin6e pour H. primigenium en haut et pour tt. mediterraneum de Pikermi en bas. MC III distal articular breadth plotted to total length ; logarithmic data. 95 % equiprobability ellipses drawn for 11. primigenium (above) and 11. mediterraneum (below) from Pikermi to facilitate comparisons. f Fig. 3 ~ Diagramme de dispersion de la largeur articulaire distale des MT 1II en fonction de la Iongueur totale (6ch. logarithmique). Ellipses probabilit6 id. fig. 2. MT II1 distal articular breadth plotted to total length ; logarithmic data. 95 % equiprobability ellipses drawn for H. primigenium (above) and 1-1. mediterraneum (below) from Pikermi to facilitate comparisons. .78 S'* K .70 .66 .42_ .48 .44 .40. / I ! K~H 1 Phal. 2 vol. I/br f Fig. 5 - Diagramme de dispersion de la largeur des phalanges I Ien fonction de leur longueur plantaire (6ch. logarithmique), id. fig. 2 Phalanx 2 breadth plotted to volar length ; logarithmic data. 95 % equiprobability ellipses drawn for H. primigenium (above) and H. mediterraneum (below) from Pikermi to facilitate comparisons. t Fig. 4 - Diagramme de dispersion des hauteurs de i'astragale en fonction de la largeur distale (6ch. logarithmique), id. fig. 2. Astragalus height plotted to distal breadth ; logarithmic data. 95 % equiprobability ellipses drawn from Pikermi to facilitate comparisons. - - 34 . . . . the teeth of H. praesulcatum, as well as of H. mediterkaneum, is described as rounded or only slightly elongated. It has repeatedly been stressed (Lungu, 1973) that in teeth of fossil horse species plication frequency and protoconal shape and size vary bet- ween individuals, between categories o,f teeth, and as a result of wear. In a sample of teeth of type H. primiEenium (von MEYER) from the Rhine-valley Forsten (in press) found protoconal length to vary between 0,5.7-1,03 cm, and plication frequency between 4-60 plications. These tooth characters, thus, have to be critically weighted. ~In species of fossil horses the teeth are seldom morphologically >, and single, abberrant specimens in a sample usually cannot, nor should, be considered specifically different from the rest of the sample. Analysis shows the material from Nesebr proba- bly to have been sampled from a single population. The skulls, e.g. PIN 130 adult and juvenile, both referred to H. nesebricum, and N ° 33 and 34 referred to H. praesulcatum, clea~ly belong in a single species (Table 1). The preorbital ,fossa is oval to pershaped and situated far in front of the eye. The characteristics of Hipparion from Nesebr agree with those of type H. primigenium (Forsten, in press) : metapodials and other lirrrb bones rela- tively massive, ahhought not much so (Table 2) ; protocone often long and narrow; plication fre~ qency rather high ; cingular structures in the lower cheek teeth variable. I believe Sarmatian hipparion in Bulgaria, although slightly smaller than the type form, to represent the wide-spread species H. primigenium. Kalimantsi The material from Kalimansti in the Struma- basin a ccordin 9 to I. Nikolov (1973b) derives from several stratigraphic levels, including Meotian to Middle Pontian. In the Meotian Nikolov identifies H. mediterraneum, H. praesulcatum, H. cf. theo- baldi LYDEKXER, and H. sp B (on p. 17 he more or- less clearly refers H. sp. B to H. nagriense HUSSAIN). In the Lower Pontian are said to occur H. mediterraneum, H. praesulcatum, H. matthewi ABEL, and H. microtatum NIKOLOV, and in the Middle Pontian H. mediterraneum, H. praesulca- turn, and H, c[. crassum GERVAIS. Hipparion cf. theobaldi, H. cf. crassum, and H. sp. B are a,ll identified on the basis of single or a few tooth rows, or isolated teeth ; to H. of. crassum Nikolov (p. 11~12) also refers a metacarpal (SLI 3'01). The same remarks as above, regarding the use of single teeth or even tooth rows in taxonomy of horse species, are valid here. The total tooth material does not appear to be as variable as 'would be expected if seven different species were involved. The large, strongly plicated teeth of H. praesul, catum, H. cf. theobaldi, H. cf. crassum, and H. sp. B, in addition to t'he large, but moderately plicated specimens which I. Nikolov refers to H. medi~erraneum, probably all belong in one species. The teeth referred to H. matthewi, on the other hand, are strongly worn specimens, in most cases belonging to a middle-sized form. The limb bones, especially the metapodials which are in majority, rather clearly indicate that only two forms of H(pparion are present (fig. 2 to 5), Moreover, these forms seem to correspond to those found at the classical site Pi,kermi. Thus there is a large, massive hipparion, not signifi- cantly different from H. primigeniurn, to whi,ch also the large teeth are referable (Table 3, 4). Equally common in this material is a smaller and rather more slender form, similar to type H. mediterra- neum at Pikermi (Table 5, 6), It must be stressed that Nikolov's H. Mediterraneum is different from H, mediterran.eum ROTH ~ WA6NEr~ at Pikermi, and rather correspond to H. brachgpus HF.~SEL = H. primigenium (v. MEYnR). Kromidovo The stratigraphic range of the material from the fossil locality Kromidovo, also in the Struma-basin, is said to be similar to that of the material from Kalimantsi. Species identified in the Meotian are H. mediterraneum and H. praesulcatum ; in the Lower Pontian H: mediterraneum, H. praesulca~ turn, H. strimoniense NIKOLOV, H. matthewi, and H. microtatum, and in the Middle Pontian H. mediterraneum and H. praesulcatum (Nikolov, 1973b). Analysis of the data, in particular data on the metapodials (fig. 1~4), indicates two forms, probably identical to those at Kalimantsi. The large and by far commonest form at this site seems slightly larger and more massive than the large form at Kalimantsi (Table 7, 8). The ~roportions of the limb bones of the small form ,from Kromidovo are not with certainty known, but appear to correspond to those of H. mediterran,eum (Table 9). I. Nikolov (p. 10-11) describes a new species, H. strimoniense, on the basis c~f a skull and jaw, and some limb bones (N ° 350/1 - 17); the material was belived to derive from a single individual. However, the skull and jaw, which are 35 those of a juvenile with M1/1 barely emerging, .clearly do not belong together with the limb bones, 'which are those of adults with all epiphyses well fused with the diaphysis. I. Nikolov especially stressed the unusual length and slenderness of the metapodial. This MT HJ ,(N ° 350/10), which heavily restored with plaster measures 28.4 cm in length, may in fact be considerably shorter, more nearly comparable to 24.5-26.0 cm for the slender metapodials from Kalimantsi. A~Ithough I. Nikolov (1973b, p. 13) mentions H. matthewi from the Pontian of both Kalimantsi and Krimidovo, it is uncertain whether dwarf- hipparion does occur at these localities. I have not seen limb bones of dwarf hi.pparions, and in all cases the teeth referred to this species are much 'worn specimens and accordingly appear to be relatively small: e.g. Kalimantsi GLI 83, GU 85, and 6 specimens without number, including skull fragments, Gorna Sus.china N ° 56 and one specimen without number. A skull (SU 5'88, Table 12) from the Pontian of Ploski Blageovradsko, described as a new species, H. microtatum, clearly does belong to H. matthewi, and a small, water- worn left M '~ (N ° 13'?) from the Meotian at Ezerovo, may also pertain to this species. The geographic rang of this small 'form, typically pre- sent in the upper beds at Samos, is thus extended. I examined samples from other Bulgarian sites as well, but less is preserved than from the main sites mentioned. These samples chiefly consists of isolated teeth. r. Nikolov (1973,b, p. 7, 10) identifies H. medi- terraneum from the Meotian to Middle Pontian of Hadjidimovo in the Strums basin, and H. prae- sutcatum from the Post-Pontian Levantine of Iva- nane and Hrabrsko in the Sofia-basin. The materia,1 from these sites derives from a large, massive form ; measurements on the teeth and limb bones correspond to those of H. primigenium (Table 10, 11). DISCUSSION I. Nikolov (1973,b p. 22-26) discussed the migrations of different Hipparion species in the Old World, and traced the influence of geogra- phically different elements in the Bulgarian faunas. He believes, for example, tooth morphology of the Bulgarian hipparions, e.g. presence of cingular structures in the lowers and details Of p.lication of the uppers, to indicate genetical relationships with the African forms. In conclusion Nikolov tought that the Bulgarian hipparions are morphologically more Similar to the Asiatic and N. American forms than to the ,European ones. The three species of Hipparion present in the Bulgarian faunas studied are in fact typical repre- sentatives of the contemporaneous European and SW Asiatic faunas. Hipparion primigenium, pos- sibly the sole representative of its genus in the Sarmatian (gallesian), was widely distributed in Europe, occurin 9 also from N. Africa through Turkey to India (Forsten, in press). I consider early Hipparion in Asia, e.g. in the Chinji and Nagri, a synonym of H. primigenium. As is to be expected, the species varied over its immense geographic range, but certain, evidently adaptative, traits characterize local ,popu,lations even geogra- phically widely spaced. In general in this species the plication frequency is high and the protocone may be elongated and flattened. The latter cha- racters, also present in Bulgarian Sarmatian H. primigenium, I. Nikolov (p. 9) particularly inter- preted as indicatin 9 American and Asiatic affinities. In N. American hipparions of roughly the same age, i.e. Burge-~Clarendon, the protocone was va- riable, but tended to be flattened in the larger forms (Webb, 1969; Forsten, 1975). Or~ly later, in the Hemphillian and Kimballian, did hipparions of the Neohipparion- grou.p sport an elongated and strongly flattened protocone. The > characters of early Hipparion in Bulgaria, thus, are characters shared with most local H. primigeniurn. This species survived in the Pikermian in Central Europe, co-habiting with other species Southern Europe and the neigh.bou- rin 9 parts of Asia. In these marginal faunas H. primigenium occurred together with more slender- built and smaller forms, e.g.H, mediterraneum and H. matthewi. H. mediterraneum was, as I. Nikolov (1973b, p. 23-24) pointed out, widely distributed on the Balkan Peninsula, in Bessaradia and the Near East; this was also the geographic range of /-/. matthewi. I. Nikolov thought that H. - - 36 rum >>, i.e. H, matthewi, shows Asiatic and N. American affinities. Dwarf hipparions occurred rather early in N. America, culminating in Blancan Nanhippus, and many Asiatic hipparions were small. In the Old World dwarf hipparions ~parti- cularly occurred in areas surrounding the Medi- terranean; the Iberian Peninsula being one center of their evolution, the eastern part of the area another. The type locality o,f H. matthewi is Samos, but the species occurred throughout Tur- key north to Bulgaria and Bessarabia. M. Pavlov (189'0) described a metapodial fragment ~from Sevastopol, the Crimea, as Hipparion ? minus, and C. Przemysiki (1912) commented on some small specimens, called H. sp. (var. minor), from the Meotian of Kuijalntk on the Black Sea W coast ; these occurrences represent H. matthewL In the northern parts of its range H. matthewi seems to have been relatively rare. The list of the bulgarian Hipparion is actually as follows : Nesebr - H. primigeniura (v. MEYER) Kalimantsi ~ H. primigenium (v. MEYER) H. mediterraneum ROTH ,~ WAGNER Kromidovo - H. primigenium (v. MEYER) H. mediterraneum ROTH ,~ WAGNER' Ploski Blagoevradsko . H. matthewi ABEL Ezerovo - H. matthewi ABEL Hadjidimovo . H. primig, enium (v. MEYER) Ivanane - H. primigenium (v. MEYER) Hrabrsko . H. primigenium (v. MEYER) The affinities of the Bulgarian hipparions are 'those to be expected in consideration of their geographic and stratigraphic situation. In fact, at any one time then existing species of Old World a) Ill O O em K @ t- O O o USS R W. EUROPE L eva ntinian Pontian ~Aeotian Sarmatian Perpignanian Pikermian Vallesian Fig. 6 - Tableau des correspondances stratigraphiques entre les formations de Russie et d'Europe occidentale. Table of stratigraphic terminology used in the text. Correspondence between Russian and W. European terms appro- ximate. hipparions were widely distributed, indicating an absence of serious barriers to their spread. It is, accordingly, rather superfluous to distinguish between > and > characteris- tics. Even the rather well defined dental charac- teristics of the Late Pliocene-Early Pleistocene African hipparions were already clearly foreboded in earlier 'forms on that continent as was as in Europe and Asia (Forsten, 1968). Manuscrit regu le 11-07-77 Manuscrit d4finitif regu le 20-11-77 - - 37 - - R E F E R E N C E S BAKALOV P. 8 NIKOLOV I. (1962). - - Fossilite na Bulgarija. X. Tertierni bozainitsi. Bolgarska Akad. Naukite, Sofia, p. 1-162. FORSTEN A. (1968). - - Revision of the Palearctic Hipparion. Acta Zool. Fennica, Helsinki, 119, p. 1-134. FORSTEN A. (1975). - - The fossil horses of the Texas Gulf Coastal Plain: a revision. The Pearce-SeUars, s. 22, p. 1-8'6. FORSTEN A. (in press). - -Hippar ion primigenium (von MEYER), an early three-toed horse. LUNGU A.N. (1973). - - Novi vid Hipparion iz srednevo sarmata Moldavii. Pal, eontologija i stratigra[ija mesokainozoa joschnich okrain russkoi platform, p. 87-115. NIKOLOV I. (1971). - - Neue Vertreter der Gattung Hipparion in Bulgarien. Bulgar. Akad. Sci., Sofia, Com. Geol., Paleontol. 20, p. 107-1221 NIKOLOV I. (1973a). - - U:ber einigen Adaptations- prozessen bei den Hipparionen. Bulgar. Akad. Sci., Sofia, Paleontol. 22, p. 71~80. NIKOLOV I. (1973b) . - Gattung Hipparion in Bulgarien. Bulgar. Akad. Wiss. Geol. Inst., Sofia, p. 1-27. PAVLOV M. (1889). - - Etudes sur l'histoire pa,l~on- tologique des ongul~s. Bull. Soc. Imp. Natur. Moscou, 3, p. 653-716. PRZEMYSKI ,C. (1912). - - Recherches pal4ontologi- ques du gisement des ossements fossiles des ter- rains m4otiques pros d'Odessa. Zan; Novoross. Obschest. ]estjestvosn., Odessa, t. 39, p. 1-13. WEBB S. D, (1969). - - The Burge and Minnecha- duza Clarendonian mammalian faunas. Univ. Cali[. PubL Geol. Sci., Berkeley, 78, p. 1-191. - - 38 - - Var iate p2. M 3 length p2 . p4 length M 1. M 3 length p2 _ orbit 11 .p2 - 2 13 . p2 snout width p2 - 2 width palatal width P2 " M3 length P2 " P4 length M 1 - M 3 length ramus depth P2 ramus depth P4 "ramus depth M 3 I3 " P2 11 " gonion snout width symphysial I O.R. N M + 14.9 + 15.0 2 7.97 - 8.55 5 8.21 + .108 + 6 .30- 6.80 3 6.53 + 16.5 17.6 2 12.5 1 9.38 1 6.22 1 6.60 7.18 2 -+ 10.7 10.85 2 14.5 - 16.7 6, 15.54 + .293 6.93 - 8.82 6 7.56 + .299 7.33 - 8.10 6 7.23 + .128 4 .75- 5.55 4 5.14 6 .04- 6.88 5 6,45 + .152 8 .55- 9.38 3 9.08 + 7.55 - + 9.40 4 +-8.60 -+ 38.0 39.0 2 4.50 5.36 2 8 .28- 9.15 3 8.69 6" .241 + .076 .719 + .207 .732 + .211 .313 + .090 .339 + .107 V 2.93 4.63 9.68 4.32 5.26 Var iate MC IIl length prox. br. MT III length prox. br. metapod, dist. radius length prox. br. tibia length dist. br. calcareum h. tuber br. Astragalus br. height Phal. 1 dors. 1 vol. 1 bredth Phalanx 2 vol. 1 Tabl. 1 - Statistics on the skull and jaw. H. primigenium, Nesebr. br. O.R. N M 20.0 - 21.6 8 20.89 + .199 3.61 - 4.18 8 3.95 + .077 22.5 - 24.7 16 23.56 + .175 4.01 - 4.26 13 4.12 + .022 3.34 - 3.90 26 3.60 + .028 25.9 1 6.84 1 34.3 1 6.51 - 7.54 6 6.81 + .161 10.57 10.77 2 4.20 4.23 2 3.93 - 4.35 7 4.16 + .069 5.33 - 5.91 6 5.60 + .099 5.30 - 5.80 3 5.52 5.32 5.43 2 2.75 - 2.90 3 2.88 3.68 - 3.87 4 3.77 6" .563 + .141 .219 + .055 .698 + .123 .08O + .016 .142 + ~019 .393 + .114 .184 + .049 .242 + .070 Tabl. 2 - Statistics on the limb bones. H. primigenium, Nesebr. V 2.69 5.54 2.96 1.95 3.96 5.77 4.28 4.32 - - 39 - - Variate p2 . M 3 length p2 . p4 length M 1 - M 3 length p2 - 2 breadth palatal width P2 " M3 length P2 " P4 length M 1 - M 3 length ramus depth P2 ramus depth P4 I3 " P2 snout width symphys. 1 Variate MC I I I length prox. br. MT III length prox. br. metapodial dis. b. Radius prox. br. dist. br. Tibia dist. br. Calcaneum height tuber br. Astragalus br. d. height Phalanx 1 dots. 1. volarl. breadth Phalanx 2 vol. 1. breadth O.R. N M 15.2 - 15.9 3 15.6 8.18 - 9.24 7 8.60 +..146 6.48 - 7.16 4 6.81 5.88 1 10.64 1 15.0 15.6 2 7.60 - 8.85 9 8.01 + .145 7.10 - 7.21 5 7.16 + .024 4.45 - 5.64 3 4.90 5.93 - 6.77 4 6.26 8.36 1 -+ 4.35 1 8.00 1 6"-" .387 + .103 .435 + .102 .055 + .017 Tabl. 3 - Statistics on the skull and jaw. H. primigenium. Kalimantsi. : O_R. N M i f" 19.6 - 21.0 3 20.3 4.30 4.38 2 4.34 23.0 - 24.5 5 23.87 + .274 4.04 4.80 i1 4.27 + .068 3.41 - 3.97 18 3.66 + .035 6.96 1 6.35 1 6.32 - 6.64 4 6.54 10.38 10.56 2 10.47 4.24 - 4.31 3 4.27 4.49 - 4.53 3 4.51 5.62 - 5.77 3 5.72 5.65 1 5.55 1 3.05 1 3.58 3.83 3 3.72 3.04 3.18 4 3.10 .612 +..194 .226 + .048 .148 + .025 Tabl. 4 - Statistics on the limb bones. H. primigenium. Kalimantsi. V 4.49 5.43 7.65 V 2.56 5.28 4.05 Variate p2 . M 3 length p2 . p4 length M 1 - M 3 length p2-2 width P2 " M3 length P2 " P4 length M 1 - M 3 length ramus depth P2 ramus depth P4 ramus depth M 3 snout width Variate MC III prox. br. MT III length prox. br. metapodial dist. Tibia dist. br. Calcaneum h. tuber br. Astragalus br. d. height Phalanx I dors. I. vol. I . breadth Phalanx 2 vol. I. breadth Variate p2 . M 3 length p2 . p4 length M 1 - M 3 length palatal width P2 " M3 length P2 " P4 length M 1 - M 3 length ramus depth P2 40 - O.R. N 13.8 14.3 2 7.38 - 7.88 4 5.72 - 6.50 6 5.43 1 14.0 - 14.6 3 7.26 - 7.60 6 6.50 - 7.03 5 5.56 1 5.47 - 6.42 3 6,97 8.50 2 5.21 1 M o'- V 7.67 6 .11+ .143 .350+ .101 14.37 7.45 + .068 .167 + .048 6.77 + .086 .192 + .061 5.79 Tabl. 5 - Statistics on the skull andjaw.H.mediterraneum, Kalimentsi. O.R. N M 3.52- 3.69 3 3.59 24.7 - 26.0 3 25.5 3 .70- 3.94 7 3.78 2 .62- 3.41 20 3.10 5.13 - 6.03 7 5.58 + 10.05 10.19 2 3 .89- 3.98 4 3.93 3 .67- 4.00 4 3.83 4 .97- 5.32 3 5.15 4.92 5,03 2 4.75 1 2.19 2.38 2 3.16 3.32 2 2.53 1 6- + .032 .084 + .022 + .042 .187 + .029 ,116 .306 + .082 Tabl. 6 - Statistics on the limb bones. H. mediterraneum, Kalimentsi. O.R. N M 15.7 1 8.46 1 6.99 6.99 2 11.24 1 15.4 1 7 .74- 8.63 3 8.08 7.35 1 4.35 4.48 2 4.42 Tabl. 7 - Statistics on the skull and jaw. H. primigenium. Kromidovo. 5.73 2.25 2.84 V 2.22 6.04 5.49 - - 41 - - Variate O.R. N M MC III length 19.5 - 21.9 3 20.37 prox. br. 4.01 - 4.46 3 4.23 MT III length 24.0 - 25 .1 5 24.57 + .224 prox. br. 4.05 - 4.64 5 4.40 + .110 metapodial dist. b. 3.64 - 4.06 14 3.80 + .035 Radius dist. br. 6.57 6.67 2 6.62 Tibia dist. br. 6.70 - 7.70 4 7.12 Calcaneum height 10.90 - 11.41 4 11.05 tuber br. 4.19 - 4.70 4 4.50 Astragalus br. d. 4.15 - 4.70 3 4.48 height 5 .76- 5.98 3 5.86 Tabl. 8 - Statistics on the limb bones. H. primigenium. Kromidovo. if-- .500 + .158 .246 + .078 .131 + .025 V 2.03 5.60 3.45 Variate metapodial dist. br. Radius length prox. br. dist. br. Tibia dist. br. Astragalus br. d. height Phalanx I dors. I. vol. I. breadth Variate MT III prox. br. metapodial dist. Radius length prox. br. dist. br. Astragalus dist. height Phalanx I dors. I vol. I breadth Phalanx 2 vol. I. breadth O.R. N M 2.95- 3.16 3 3.05 _ 26.4 1 6.12 1 5.66 1 6.20 1 3.53 1 4.90 1 5.24 5.64 2 5.44 4.98 5.25 2 5.12 2.25 2.62 2 2.44 Tabl. 9 - Statistics on the limb bones. 11. mediterraneum. Kromidovo. O.R. N M 4.88 1 3.81 4.07 2 3.94 27.4 1 7.32 1 6.76 1 4 .24- 4.41 3 4.35 5 .51- 5.81 3 5.71 5.96 1 5.73 1 2.91 1 3.91 1 3.27 1 TaN. 10 - Statistics on the limb bones. 11. primigenium. Hadjidimovo. - 42 - Var ia te MC III length prox. br. MT III length prox. br. metapodial dist. Radius length dist. br. Calcaneum height tuber br. Astragalus br. d. height Phalanx I dots. I vol. I. breadth Phalanx 2 vol. 1 breath O.R. N 20.6 20.7 2 4.11 4.32 2 23.0 1 4 .29 1 3.62 - 3.91 3 27.7 28.2 2 6.28 6.46 2 10.67 10.73 2 4.06 4.09 2 4.31 4.48 2 5.67 5.69 2 6.11 1 5.94 1 3.06 1 3 .94- 4.02 3 3 .27- 3.47 3 Tab1. 11 - Statistics on the limb bones. Variate O.R. N p2. M 3 length 12.54 1 p2 . p4 length 6.90 1 M 1 - M 3 length 5.55 1 p2 .orb i t + 13.1 1 p2-2 width 5.65 1 palatal width + 8.88 1 M 3.76 3.98 3.38 H. primigenium. Ivanane. Tabl. 12 - Statistics on the skull. It. matthewi. Ploski Blagoevradsko.