During the Pleistocene, there were three main groups of Im Pleistozän traten drei Hauptgruppen von Nashörnern auf, rhinoceroses, each of them in a different part of the Old jede in einem anderen Teil der Alten Welt: die afrikanische World: the African lineage leads to the modern square- Linie führt zu den heutigen Breitmaul- und Spitzmaulnashör- lipped rhinoceros and black rhinoceros, the Asian group nern, die asiatische Gruppe umfasst das Panzer-, das Suma- includes the great one-horned rhinoceros, the Sumatra tra- und das Javanashorn sowie ihre Vorfahren. Zur dritten rhinoceros and the Java rhinoceros as well as their ances- Gruppe, die im späten Pleistozän ausstarb, gehören Coelo- tors. The third group, which became extinct in the Late donta und Stephanorhinus. Das Wollhaarnashorn (Coelodonta Pleistocene, includes Coelodonta and Stephanorhinus. The antiquitatis) trat in Europa zum ersten Mal während der woolly rhinoceros (Coelodonta antiquitatis) appeared in Elsterkaltzeit auf. Stephanorhinus kirchbergensis, das Wald- Europe for the first time during the Elsterian cold period. nashorn, ist auf die Interglaziale beschränkt und wanderte Stephanorhinus kirchbergensis, the forest rhinoceros, is lim- wahrscheinlich nach jeder Kaltzeit erneut von Asien aus ein. ited to the interglacial periods and probably dispersed again Das Steppennashorn (Stephanorhinus hemitoechus) ist wie- and again after each cold period from Asia into Europe. The derum in Europa seit 450 000 Jahren heimisch. In Neumark- steppe rhinoceros (Stephanorhinus hemitoechus) again has Nord konnten diese drei Nashörner zusammen nachgewiesen been present in Europe for 450,000 years. All three types werden, was umso bemerkenswerter ist, weil das Wollhaar- of rhinoceros together could be documented in Neumark- nashorn im Allgemeinen als Vertreter der Glazialfaunen gilt. Nord, which is even more astonishing, as the woolly rhinoc- Als Besonderheit wurden aus den Backenzähnen der Nas- eros generally is considered to belong to the glacial fauna. hörner von Neumark-Nord noch pflanzliche Nahrungsreste Remarkably, from the molars of the rhinoceros from Neu- geborgen. Als Nahrung der Waldnashörner konnten Reste mark-Nord plant remains were recovered. It could be proven von Birken gewächsen, Rosengewächsen und Gräsern sowie that the food of the forest rhinoceros included Betulaceae, insbesondere von Pappeln, Eichen, Weißdornen, Feuerdornen, Rosaceae and grasses and, in particular, from poplars, oaks, Brennnesseln und Seerosen festgestellt werden. Bedauer- hawthorns, pyracanthas, stinging nettles and water lilies. licherweise war es nicht möglich, vom Steppennashorn und Unfortunately, no food remains could be identified from the vom Wollhaarnashorn Nahrungsreste zu identifizieren. steppe rhinoceros and the woolly rhinoceros. The rhinoceroses from Neumark-Nord and their nutrition DIE Nashörner VON NEUmarK-NOrd Und Ihre ErnÄhrUNG Jan van der Made and René Grube Rhinoceroses are mammals that share a number of a land connection, many kinds of mammals dispersed characters, such as hoofs, and have an odd number of from Asia into Europe, including the rhinoceroses. This them per hand or foot, as well as cheek teeth with a is an important event, called the Grande Coupure. In typical pattern of crests. Contrary to what one might a similar way, the Middle East started to become land, expect, horns were not present in many species of rhi- intermittently connecting Africa and Eurasia, allowing noceroses. during the Early Miocene, around 2o Ma ago, the rhi- The family Rhinocerotidae originated in the Mid- noceroses to reach Africa and little later proboscideans dle Eocene of North America and spread by the Late to reach Europe. Rhinoceroses never reached South Eocene to Asia (Cerdeño 1998). At that time Europe was America, or Australia. Rhinoceros diversity reached isolated from Asia by a sea way. When, at the beginning a peak during the Miocene. The family went extinct of the Oligocene, some 33 million years ago, tectonics in North America at the beginning of the Pliocene, and sea level fluctu ations caused the establishment of about 5 million years ago. In Europe diversity started // Die Nashörner von Neumark-Nord und ihre Ernährung 383 e st il u 1 c a s r si g s i n s s The stratigraphic distribution of s rob n e hu hu ime g c c e the European Pleistocene rhi- s a e 18 e er h u o δ0 (‰) o nt s t c t i i o s noceroses. hb c nd m m r od i l k he hu etru he e Age in millions of years, palaeo- . Co S. S S. S 3.0 S. magentism and Oxygen Isotope 5.5 5.0 4.5 4.0 3.5 Stages on the left; temporal dis- 0.0 1 La Ventana tribution of the rhinos indicated Arroyo del Culebro 2 3 Abric Romaní with black lines in the central Can Rubau 4 Crayford part of the figure; localities on Burgtonna 5 Taubach the right. Horizontal bars indi- Barrington cate presence of a species in a Schweinskopf 6 Wannen locality. Blue indicates presuma- NeumarkNord bly glacial and red interglacial Ehringsdorf 7 La Fage localities. Question marks in Ariendorf 1 front of a locality name indicate 8 Steinheim pS Steinheim aS uncertainty about the strati- Orgnac3 graphic position and a question 9 ? Daxlanden? Heppenloch mark behind a name indicates 10 ArtenacIV-V uncertainty about the taxonomic ? Kirchberg 11 Bilzingsleben identification. The figure differs from earlier versions in that 12 AragoCM III Mosbach3 Mosbach 2 is placed one cli- Mosbach2 0.5 13 Hundsheim matic cycle higher, because of Mauer+ Boxgrove ? Jockgrim doubts on the differences ? Bammental 14 between deer from Mauer and Mosbach 2 and because of the 15 Soleilhac Isernia recognition of assemblages with Süßenborn Equus suessenbornensis and 16 BadFrankenhausen Arvicola, which should be older 17 Voigtstedt WestRunton than Mosbach and younger 18 than Süßenborn. AtapuercaTD8 Die stratigraphische Verteilung 19 AtapuercaTD7 der Europäischen Nashörner aus 20 AtapuercaTD4-6 dem Pleistozän. 21 DornDürkheim3 Jahresangaben in Millionen 22 Huescar Jahren, Paläomagnetismus und 23 Sauerstoffisotopenstufen links, 25 die zeitliche Verteilung der Nas- 26 hörner ist mit schwarzen Linien 1.0 Untermaßfeld in der Mitte der Darstellung ange- Vallonnet zeigt, Fundorte auf der rechten 30 Seite. Horizontale Linien geben 31 das Vorkommen einer Spezies an einem Ort an. Blau markierte 34 Fundorte sind vermutlich glazia- 35 len, rot markierte interglazialen 36 Pirro Nord? Charakters. Fragezeichen vor 37 Venta Micena einem Ortsnamen bedeuten Unsi- cherheit in der stratigraphischen Lage und ein Fragezeichen hinter einem Namen bedeutet Unsi- cherheit in der taxonomischen Identifikation. Die Darstellung unterscheidet sich von früheren Versionen dadurch, dass Mos- to decline after the early Late Miocene (Heissig 1999) Africa, southern Asia, and northern Eurasia. The Afri- bach 2 aufgrund von Zweifeln an and they went extinct around the Pleisto-Holocene can rhinos are the lineages leading to the living black den Unterschieden zwischen den transition. and white rhinos (Diceros bicornis and Ceratotherium Rehen von Mauer und Mosbach 2 sowie bekannt gewordener Ver- Rhinoceroses tend to be large and some species simum, respectively). These species are characterized gesellschaftungen mit Equus are among the largest land mammals that existed, by the absence of incisors and the presence of two well suessenbornensis und Arvicola, with maximum body weights of up to 15–2o tonnes developed horns and massive nasals; a »cauliflower- die älter als Mosbach und jünger als Süßenborn sein dürften, einen (Fortelius/Kappelman 1993). structure« marks the position of the horns. The south klimatischen Zyklus höher Three major groups of rhinos occurred in the Pleis- Asian rhinos include the Indian, Javan and Sumatran platziert ist. tocene, each one in a different part of the Old World: rhinoceroses (Rhinoceros unicornis, R. sondaicus, and 384 Jan van der Made und René Grube // The rhinoceroses from Neumark-Nord and their nutrition Dicerorhinus sumatrensis, respectively) and their ances- tors. These species have large second lower incisors, which are kept sharp by wear against small upper inci- sors. The incisors are used in fighting, while the horns (one in Rhinoceros, two in Dicerorhinus) are small and a less important in fighting. The nasals are relatively gracile. The third group consists of various species of Coelodonta and Stephanorhinus and went extinct dur- ing the Late Pleistocene. Like the African rhinos, they lacked incisors and had two well developed horns, but 2 differed from those in having elongate nasals supported b Left view of the skulls of the by an ossified nasal septum. The position of the horns rhinoceroses from Neumark- Nord: a) no. 198 – Stephanorhi- is again indicated by »cauliflower-structure« on the nus kirchbergensis; b) nasals. The horns are made of keratin (the same mate- HK88:14’3 – Stephanorhinus rial as nails and hair), which usually does not fossilize, hemitoechus; c) 1996, 47 – Coelodonta antiquitatis. but fossil horns of Coelodonta are known and are very c Linke Ansicht der Nashorn schädel long. There is a further rhinoceros in the Pleistocene of von Neumark-Nord: a) Nr. 198 – northern Eurasia, called Elasmotherium, which is more Stephanorhinus kirchbergensis; b) HK88:14’3 – Stephanorhinus distantly related to the three major groups of rhinos. hemitoechus; c) 1996, 47 – It is a very large form with a huge dome in the mid- 10 cm Coelodonta antiquitatis. dle of the skull, which is commonly assumed to have supported a huge horn. It lived in Asia and Eastern Europe, though there are some problematic records in woolly rhino (Coelodonta) appeared in Europe. The Western Europe, possibly of Middle Pleistocene age scientific name refers to a structure in the upper teeth. (Guérin 198o; Cerdeño 1998). This genus is known already from the Late Pliocene of northern China and is typical of open or dry land- THE rhinoceroses of the Pleistocene of EuroPE scapes. During glacial periods such landscapes spread The European species of Pleistocene rhinoceros belong into Europe and so did the woolly rhino. Initially it to the third group. Figure 1 gives their distribution in reached Germany, later it appeared in France and dur- time.