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The Oldest Eurasian Hominoid

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The Oldest Eurasian Hominoid Elmar P. J. The oldest Eurasian hominoid Heizmann Staatliches Museum fu¨r Engelswies is an early Miocene vertebrate locality in southern Naturkunde, Rosenstein 1, Germany with a rich assemblage of terrestrial mammals, invertebrates D-70191 Stuttgart, Germany and fossil plants. It is dated to 16·5–17·0 Ma based on magneto- stratigraphy, biostratigraphy and lithostratigraphy, and includes David R. Begun among the faunal remains a hominoid upper molar fragment, the Department of Anthropology, oldest hominoid so far identified from Europe. The evidence from University of Toronto, Engelswies suggests that hominoids arrived in Eurasia about 17 Ma, Toronto, Ontario M5S 3G3, roughly contemporaneously with pliopithecoids and Deinotherium, Canada and before the last marine transgression to isolate Eurasia from Africa. Thick enamel and low dentine penetrance may have been key Received 6 February 2001 adaptations that contributed to the success of hominoids of dentally Revision received 5 June modern aspect in western Eurasia and ultimately to their ability to 2001 and accepted 7 June spread to eastern Eurasia and Africa in the middle and late Miocene. 2001 2001 Academic Press Keywords: early Miocene, thickly enameled hominoid, Eurasia, paleobiogeography, Journal of Human Evolution (2001) 41, 463–481 dispersal events. doi:10.1006/jhev.2001.0495 Available online at http://www.idealibrary.com on Introduction Engelswies, which has been quarried for limestone since the nineteenth century. The Tertiary fossil localities in southwest quarry has long since been abandoned, and Germany are numerous and critical to cur- the site is covered in dense vegetation (the rent interpretations of Cenozoic biostratig- Talsberg is protected today as a natural raphy, biochronology and paleoecology. monument). Several fossil localities Most of the Neogene localities in this region (Engelswies 1–4) are known from quarry are associated with erosional and deposi- activities in the Talsberg and have been the tional regimes linked to the Alpine Orogeny subject of research since the late nineteenth and global climate changes that together century. It is particularly well known for its resulted in periodic opening and closing of beautifully preserved plant remains and the seaway between the Atlantic and Indian invertebrates, which include freshwater Oceans. Many of these localities preserve a crabs and gastropods (Quenstedt, 1885; rich record of flora and fauna from the past, Engel, 1908; Fraas, 1910; Kla¨hn, 1922, including the famous sites of Langenau, 1924, 1925, 1926; Rutte, 1953; Pretzmann, Steinheim and Ho¨wenegg, all of which con- 1987; Schweigert et al., 1997). tain many nearly complete skeletons of ter- Neglected for many years, Engelswies was restrial fossil mammals (Heizmann, 1992). the subject of renewed excavations by Heinz Among these localities is Engelswies, near Tobien (Mainz University) in the 1970s, Sigmaringen, south of Stuttgart and east of during which the tooth described here was Freiburg (Figure 1). The locality is situated found. Tobien found the tooth, which he in the Talsberg, southwest of the town of recognized as hominoid but never described, Address correspondence to: David R. Begun, on 24 June 1973, in the lower part of the Department of Anthropology, University of Toronto, Toronto, ON M5S 3G3, Canada. E-mail: begun@ profile of the Scherer quarry (excavation chass.utoronto.ca horizon 4 in Figure 2), from which most of 0047–2484/01/100463+19$35.00/0 2001 Academic Press 464 . . Figure 1. Geologic map with the locality of Engelswies (arrow) and other important localities, major rivers and large cities in the region. Inset shows the location of Engelswies in Western Europe. 465 Schichten (Figure 2). From these cores there is no evidence of any disconformity, indicating a continuous record of sedimen- tation. This, and the faunal content of the fossiliferous layers, indicates that these sedi- ments correlate to the top of the Kirchberger Schichten, the basal facies of the OSM [Obere Su¨ßwassermolasse (or Upper Freshwater Molasse in English)]. Figure 2. Geologic section and stratigraphy of the The molasse basin sediments of southern Engelswies locality. The hominoid tooth is from hori- Germany and Switzerland are among the zon 4. Stratigraphic column modified from Ziegler best studied of any Tertiary sediments in (1995; Abb. 2). Geologic section modified from Schweigert, 1992; Abb. 2). Legend: (a) oncolite (free Europe, and the sequence of lower and rolling or mobile structures of algal origin) facies, (b) upper marine and freshwater layers and oncoidal crusts and plant fragments, (c) limestones numerous marker horizons, including the with terrestrial detritus, (d) lake sediments. Abbrevia- tions: n=north, s=south, osm=upper Freshwater Kirchberger Schichten, are well established Molasse, ki=Kirschenberger Schlichten, usm=lower (Gall et al., 1977; Reichenbacher, 1989; Freshwater Molasse, q=Quaternary overburden, Schlunegger et al., 1996; Bolliger, 1997; j=Jurassic limestone. Heissig, 1997; Kempf et al., 1997; Reichenbacher et al., 1998; Kempf & the large mammals are known (Schweigert, Matter, 1999; Sach, 1999)(Figure 3). It has 1992). This horizon is 4 m below the level been demonstrated that the Kirchberger yielding the majority of the micromammals Schichten marks the transition between the on which the biostratigraphic correlation is upper marine molasse sediments (OMM) mostly based (Ziegler, 1995). In 1990, and the upper Freshwater Molasse Tobien deposited the tooth in the Staat- sediments (OSM) (Heizmann et al., 1980; liches Museum fu¨r Naturkunde in Stuttgart Heizmann, 1992; Schweigert, 1992; (SMNS) along with the other mammalian Reichenbacher, 1993; Ziegler, 1995; fossils from Engelswies. This specimen has Reichenbacher et al., 1998). This transition been noted in the literature (Heizmann, occurs before the onset of the Langhian 1992; Andrews et al., 1996; Heizmann et al., transgression, a stage in the Mediterranean 1996) but never described in detail, nor marine sequence marked by the brief recon- have its implications for Miocene hominoid nection of the Atlantic–IndoPacific oceanic biogeography been fully discussed. seaway (Ro¨gl, 1999). The Langhian is well dated by magnetostratigraphic, radiometric, marine biostratigraphic and lithostrati- Geology, biostratigraphy and graphic evidence to between 16·5 and paleoecology 14·9Ma(Steininger, 1999). At Engelswies The freshwater limestones of Engelswies the occurrence of Megacricetodon bavaricus represent an isolated outcrop of an originally and the evolutionary stage of Democricetodon widespread sedimentary stratum covering mutilus allow a precise correlation to the the northern rim of the Alpine Molasse lower part of MN 5 or late Orleanian Basin. Hahn (1968) in his detailed analysis (Ziegler, 1995), which we place in the early of the regional geology took core samples at Miocene. There is currently a debate on the Engelswies and demonstrated the direct definition of the boundary between early superposition of the fossil levels on the dark and middle Miocene. We follow the pro- clays of the lower Miocene Kirchberger posal of Fahlbusch (1981), placing the limit 466 . . 467 between MN 5 and MN 6. Steininger The lithostratigraphic marker horizons of (1999), in contrast, sets the limit within MN significance to a determination of the age 5. Either way, Engelswies, in the early part of Engelswies are the OMM, OSM, of MN 5, is in the early Miocene. The Kirchberger Schichten and the Ries impact composition of the large mammal fauna is horizon. Based on the marine microfossil consistent with the biostratigraphy of the record and recent magnetostratigraphic, small mammals (Table 1). A study of lithostratigraphic and radiometric work the Anchitherium (Abusch-Siewert, 1983) has ages of these horizons in Central Europe suggested an age of MN 6 or older. The appear reasonably well established. Given suids also imply an age of MN 5. Hyotherium the ages of these marker horizons, the MN 5 soemmeringi is a typical species for MN 5, localities of Langenmoosen, Engelswies, and and the genus Bunolistriodon is extinct at the Puttenhausen are dated to between about 15 end of this zone. To summarize, based on and 16·8Ma(Steininger, 1999)(Figure 3). regional geology, lithostratigraphy from core However, in the Calatayud-Daroca Basin of samples, and mammalian biostratigraphy Spain, magnetostratigraphic and biostrati- and biochronology, Engelswies is between graphic evidence suggests that MN 5 is the base of the Langhian and the base of about 1 Ma younger (16–13·8 Ma, Daams MN 5, or 16·5to17Ma(Ziegler, 1995; et al., 1999). There are several possible Steininger, 1999)(Figure 31). resolutions to these divergent interpreta- tions. The evidence from either Central 1An important debate in European Miocene chron- Europe or Spain may be correct and the ostratigraphy currently centers on the age of the late early Miocene and middle Miocene MN zones (MN other misleading, for a number of technical 4–7/8). Evidence from Spain is interpreted to suggest reasons, or, there may be strong diachrony an age for these zones 1 to 2 Ma younger than ages between Central Europe and Spain. The derived from evidence from elsewhere in Europe (Steininger, 1999; Daams et al., 1999). For a variety of issue of the age of MN 5 is unresolved, and reasons beyond the scope of this paper, including the is in fact the focus of much attention today. fact that Engelswies predates the well dated Langhian Whatever the outcome, mammal localities
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