ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Carolinea - Beiträge zur naturkundlichen Forschung in Südwestdeutschland Jahr/Year: 2000 Band/Volume: 58 Autor(en)/Author(s): Bernor Raymond L., Fessaha Nardos Artikel/Article: Evolution of Late Miocene Hungarian Suinae (Artiodactyla, Suidae) 83-92 ©Staatl. Mus. f. Naturkde Karlsruhe & Naturwiss. Ver. Karlsruhe e.V.; download unter www.zobodat.at carolinea, 58 (2000): 83-92, 4 Abb.; Karlsruhe, 23.05.2000 83 Raym o nd L. B e r n o r & N a r d o s F essa ha Evolution of Late Miocene Hungarian Suinae (Artiodactyla, Suidae) Abstract Washington D.C. 20059; e-mail: [email protected] Neogene suids have been collected for over 100 years from and [email protected] . Hungary, yet, until recently, there has been very little publis­ hed about them. This contribution presents our current know­ ledge of Hungarian late Miocene Suinae and their evolutio­ Introduction nary, chronologic and biogeographic relationships with other Eurasian suines. We analyse an evolutionary series of Mioce­ Despite more than 100 years collection of fossil suines ne Suidae including Hyotherium meissneri, Hyotherium soem- meringi, Propotamochoerus palaeochoerus, Propotamochoe- from late Miocene Hungarian strata, very little is kno­ rUS sp. (Spain), Microstonyx erymanthius/major and H ip p o p o - wn about the actual taxa represented or the material tamodon antiquus in order to decipher evolutionary trends wit­ that it is based upon. A recent synthesis of Western hin the Suinae. Comparing bivariate plots and log-ratio plots Eurasian Suoidea by Fortelius et al. (1996) reported for all cheek teeth of this series of suids we find that suines the following fossil suines from Hungary: Propota­ exhibit serially progressively longer M1/m1, M2/m2 and mochoerus provincialis Gervais , 1859 (possible oc­ M3/m3 than hyotherines. Furthermore, Hippopotamodon currence at MN13? locality of Hatvan), Microstonyx would appear to be particularly distinct in its very large p4 major Gervais , 1848-52 (MN13 locality of Polgardi) length and width measurements. We find that Propotamocho­ erus and Microstonyx are distinct lineages that overlap In both and Microstonyx erymanthius Roth & W agner , 1854 time and space. Propotamochoerus would appear to be an in­ (MN11 locality of Csakvar). A recent study of the Ru­ dex for warm equitable forests (typical for the late Astaraci- dabanya suid fauna includes the MN8-9 suine Propo­ an/Vallesian of Middle Europe), while Microstonyx is an index tamochoerus ( = Korynochoerus of Schmidt-Kittler , for more seasonal open country woodlands typical of late Mio­ 1971 and Hellmund , 1995) palaeochoerus Kaup , cene "Pikermian” faunas. 1833 (late MN9 locality of Rudabanya; Fortelius et al.1999 and in press). These reported taxonomic re­ Kurzfassung Evolution ungarischer Suinae im späten Miozän (Artio­ cords differ slightly from those listed in the curatorial dactyla, Suidae) records of the Hungarian Geological Survey (HGI or Seit über 100 Jahren wurden in Ungarn neogene Suiden ge­ MAFI) which include: "Hyotherium" palaeochoerus sammelt, doch gibt es darüber bis heute nur wenige Veröffent­ from Rudabanya (Kretzoi et al. 1976); Microstonyx cf. lichungen. Dieser Beitrag zeigt den derzeitigen Stand der antiquus from Csakvar (Kretzoi 1954); Microstonyx Kenntnisse über die obermiozänen Suiden Ungarns und ihre erymanthius from Polgardi (Kretzoi 1954) and ano­ evolutionären, chronologischen und biogeographischen Ver­ ther latest Miocene locality, Baltavar (Kormos 1914). hältnisse zu anderen eurasischen Suiden auf. Wir analysieren die evolutionäre Reihe von miozänen Suiden, einschließlich The aim of this paper is to conduct a quantitative com­ Hyotherium meissneri, Hyotherium soemmeringi, Propotamo- parison of the Hungarian material with other pertinent cheorus palaeocheorus, Propotamochoerus sp. (Spanien), Miocene hyotherine and suine data to exact greater Microstonyx erymanthicus/major und Hippopotamodon an­ clarity in the evolution of Hungarian late Miocene sui­ tiquus, um die evolutionären Trends innerhalb der Suinae her­ nes. The Karlsruhe collections of late Miocene (medial auszuarbeiten. Durch Vergleiche von Bivarianz-Plots und Log- Turolian) Microstonyx erymanthius from Mamutghazi, Ratio-Plots für alle Backenzähne dieser Suidenserie zeigen hitherto unpublished, serve as an important basis for wir, daß die Suinae progressiv längere M1/m1, M2/m2 und our comparison and analysis. M3/m3 aufweisen als die Hyotherinae. Darüberhinaus fällt H ip ­ popotamodon durch einen sehr großen und breiten p4 auf. Wir zeigen außerdem, daß Propotamochoerus und Microstonyx zwei getrennte Linien sind, die einander in Raum und Zeit Materials and Methods überlappen. Propotamochoerus scheint demnach ein Indikator für gemäßigt warme Wälder zu sein (typisch für das späte Methods Astaracian/Vallesian Mitteleuropas), während Microstonyx als We use both continuous and discrete variables here to analy­ Indikator für eher durch Jahreszeiten beeinflußte offene Wald­ se the Hungarian suine sample, which we compare to a series gebiete ist, typisch für die obermiozänen "Pikermian"-Faunen. of hyotherine and suine pigs from Europe. The continuous va­ riables used follow the conventions set forth in Table 1 (Le­ Authors gend). Our statistical analysis here includes calculation of bi­ Prof. Raymond L. Bernor, PhD & Nardos Fessaha , College variate plots of basal length versus maximum width for all of Medicine, Department of Anatomy, Laboratory of available cheek teeth and the use of log-ratio plots of cheek Evolutionary Biology, Ploward University, 520 W St. N.W., tooth length between taxa. We use the extensive sample (over ©Staatl. Mus. f. Naturkde Karlsruhe & Naturwiss. Ver. Karlsruhe e.V.; download unter www.zobodat.at 84 carolinea, 58 (2000) 400 specimens) of Rudabánya Propotamochoerus palaeocho- by relatively short P2-3; P4 with similar length to P. pa­ e ru s as a standard for the log plots. We recognize maxillary laeochoerus-, molars exhibiting progressive serial in­ teeth as uppercase (i.e. P2, M3, etc.) and mandibular teeth as crease in length compared with hyotheres and P. pala- lower case (i.e. p2, m3, etc.). oechoerus. Hippopotamodon antiquus - A very large suine with Materials approximately twice the body mass of Microstonyx We compare the Hungarian suine species here (Table erymanthius/major/, and apparently closely related to 1 here; excluding Rudabánya specimens reported in the Microstonyx clade; characterized by p1 being pri­ Fortelius et al., in press) to other populations of Cen­ mitively long and p3 p4 longer larger and higher tral European hyotherine and suine taxa. These taxa crowned than in Microstonyx. are briefly characterized here. Hvotherium meissneri - A well known, primitive hyo­ therine typical for the latest Oligocene - basal Miocene Statistical Analysis (MN1-2) of Europe, which we have sampled from Ulm- Westtangente, Germany (Hellmund 1991). This spe­ Figures 1 a-f present bivariate analyses of maximum cies is characterized by small size, P1/p1 much smal­ width versus basal length of P2/p2 - P4/p4. P1/p1 are ler and lower crowned than P2/p2; P3 with tall pointed not plotted because of the scarcity of material, particu­ principal cusp and well defined posterolingual shelf; larly in Microstonyx species which either lack or rarely P4 with two distinct buccal cusps, a single lingual cusp preserve these teeth. P2 and p2 exhibit overlap of H. and virtually continuous cingulum; M1-M3 with small, soemmeringi and P. palaeochoerus. The lower p2 ex­ low bunodont cusps and cingulum on all but lingual hibits overlap between these taxa and both the Dorn- margin; M3 with a short talon; p4 has a principal cusp Durkheim and Turkish (Mamutghazi) Microstonyx that is moderately bifurcated lingually; mandibular mo­ erymanthius, and one specimen of Hippopotamodon lars with cingulum variably expressed on the buccal antiquus. Clear outliers in the lower p2 cluster are the surface; m1-m2 rectangular shaped; m3 an uncommon species of P. n. sp. from Spain and the Ty­ elongate/triangular shape with decreased width from pe material of Hippopotamodon antiquus from Eppels- anterior to posterior aspects of the tooth. heim. Hvotherium soemmeringi - A well represented species Figures 1c and d again exhibit more extensive overlap of hyotherine pig which is larger than H. meissneri. in the p3’s than the P3’s. There is limited overlap bet­ Our comparative sample is derived from the late early ween the Rudabanya sample of P. palaeochoerus and Miocene (MN5) of Sandelzhausen, Germany (re: the Microstonyx sample of upper P3. Mandibular p3 Schmidt-Kittler 1971). This species is characterized plots Hippopotamodon antiquus and P. n. sp. well out­ by its relatively longer premolars than H. meissneri; P3 side cluster of other plotted points. Figures 1e and 1f has a distinct posterolingual cusp; P4 cusps are es­ of upper P4 and lower p4 exhibit considerable overlap sentially as in H. meissneri, but the crown outline is of plotted points with, once again, H. antiquus plotting more quadrangular in shape and with the cingulum as a separate cluster for lower p4. less continuous on the buccal and lingual surfaces; p4 Figure 2a-f presents bivariate analyses of width versus with a more distinctly developed, lingually situated in- basal length of M1/m1 - M3/m3. Maxillary M1 exhibits nenheugel; molars generally somewhat more elongate overlap between the two hyotherine