The West Runton Mammoth (Mammuthus Trogontherii) and Its Evolutionary Signiﬁcance

Quaternary International 228 (2010) 180e209

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Quaternary International

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The West Runton mammoth (Mammuthus trogontherii) and its evolutionary signiﬁcance

Adrian M. Lister a,*, Anthony J. Stuart b a Department of Palaeontology, The Natural History Museum, London SW7 5BD, UK b School of Biological and Biomedical Sciences, Durham University, South Road, Durham DH1 3LE, UK article info abstract

Article history: The West Runton mammoth (WRM) is one of the best-stratiﬁed, most complete and best-preserved Available online 7 August 2010 skeletons of the so-called ‘steppe mammoth’, Mammuthus trogontherii. The skeleton was a male of estimated age 41 years at death. Its estimated live shoulder height of 3.9 m, and body mass of around 9 tonnes, are typical for males of the species. Together with other European skeletons, the WRM indicates a similar body size for Mammuthus meridionalis and M. trogontherii. The WRM, dating from the early part of the early Middle Pleistocene (c. 700 ka), represents an important marker in European mammoth evolution. In its mandible and dentition it has a clearly ‘advanced’ morphology overall, similar to most M. trogontherii and Mammuthus primigenius, and unlike most M. meridionalis. In its molar morphology in particular (especially plate number), the WRM already falls in the upper end of the range seen in later samples of M. trogontherii, such as those from Süssenborn and Mosbach. This early appearance of a dentally advanced mammoth runs counter to the gradualistic model of European mammoth evolution but is consistent with immigration of M. trogontherii from Asia. The WRM is more advanced than the meridionalis/trogontherii ‘mosaic’ mammoths from Voigtstedt, but previously-collected molars from West Runton hint at possible meridionalis or mosaic individuals there too. Ó 2010 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction accepted model for Quaternary mammoth evolution in Eurasia, although it is becoming clear that this was a complex geographical The mammoth lineage (Mammuthus) has provided one of the process (Lister et al., 2005). A number of largely complete skeletons most detailed examples of evolution and dispersal in the Quater- of M. meridionalis (the ‘southern elephant’ or ‘ancestral mammoth’) nary mammalian record (Lister, 1996a; Lister et al., 2005). The are known, mainly from Europe (Gaudry, 1893; Maccagno, 1962; discovery of the West Runton mammoth in December 1990 and its Garutt, 1964), and many of M. primigenius (e.g. Zalensky, 1903; subsequent excavation in 1991e5(Stuart and Lister, 2010a) yielded Garutt, 1964; Mashchenko, 2002; Lister, 2009); but of the inter- a skeleton of considerable importance to an understanding of mediate species M. trogontherii relatively few skeletons have been mammoth evolution; ﬁrst, because it is one of very few largely satisfactorily identiﬁed and described (Table 1). Until recently, complete specimens of the species Mammuthus trogontherii; and these comprised a skeleton from the northern shores of the Black second, because it was preserved within the type deposit of the Sea at Azov, southern Russia (Baigusheva and Garutt, 1987) and two Cromerian interglacial at West Runton, Norfolk, UK, with abundant individuals from the Ukraine, at Novogeorgievsk (Zakrevska, 1935) associated environmental, geological and dating evidence. and Odessa (Yatsko, 1948). The Edersleben skeleton, Germany The sequence of species Mammuthus meridionaliseM. trogon- (Garutt and Nikolskaya, 1988), belongs with the group of European theriieMammuthus primigenius (e.g. Adam, 1961) remains a widely Mammuthus transitional between M. meridionalis and M. trogontherii, but is dentally and cranially most similar to the latter (van Essen, pers. comm., 2009). From the late Middle Pleistocene, x x a skeleton from Steinheim, Germany (Dietrich, 1912) was formerly Abbreviations: R, right; L, left; dPx/dP , lower/upper deciduous molar x; Mx/M , lower/upper permanent molar x; T, thoracic vertebra; m-l, medio-lateral; a-p, referred to M. primigenius but in dentition and size is more similar antero-posterior; d-v, dorso-ventral; NCM, Norwich Castle Museum; SMG, Sedg- to M. trogontherii (Lister, 2001). A cranium with partial skeleton wick Museum of Geology, Cambridge; NMAS, Norfolk Museums and Archaeological from Gelsenkirchen, Germany, can be referred to this species but is Service; WRM, West Runton Mammoth; WRFB, West Runton Freshwater Bed. * Corresponding author. partly reconstructed (Siegfried, 1956) so is of limited research value. E-mail address: [email protected] (A.M. Lister). Dentally, the species is very well-characterised, from large samples

Table 1 Skeletons referable to M. trogontherii, arranged by year of discovery.

Locality Date of Country M3 dental Skeletal Sex Geological age Reference discovery formula shoulder (where known) height (cm) Steinheim 1910 Germany x20xex22x 370 Male 400 ka Dietrich (1912) Odessa 1911 Ukraine 20c 370 Male Tiraspolian Yatsko (1948) Novogeorgievsk 1916 Ukraine 21c 305e Femalea Zakrevska (1935) 330e (S) Gelsenkirchen 1926 Germany x19x 351e Male Late Middle Pleistocene Siegfried (1956) Edersleben b 1930 Germany x15x 345 Female Early part of early Mid. Pleist Garutt and Nikolskaya (1988) (ca. 700 ka) Azov I 1964 Russia x20x - x21x 392e Male Baigusheva and Garutt (1987); 400e (B) Bajgusheva and Titov (2007); 400e (S) Shpansky et al. (2008a,b) West Runton 1990 UK x21x 369 e Male Early part of eMPl (c.700 ka) This paper Irtysh, Chembakchino 1993 Russia x20x 350a,d Malea 550e600 ka (TL) Kosintsev et al. (2004); A. Rezvyi, pers. comm. (2009) Azov II 1999 Russia x20x - x21x 353e Female Bajgusheva and Titov (2007) 360e (B) 385e (S) Irtysh, Pyatiryzhsk 2002 Russia N12x 393e Male Tabolian Shpansky et al. (2008a,b) (est x18x) 420e (S) Kostolac 2009 Serbia N16x 374e Male Dimitrijevic, Markovic et al., in prep.

a Female gender of Novogeorgievsk skeleton determined from pelvic aperture ratio of 3.35, measured from photo in Zakrevska (1935, Fig. 37) (cf. Fig. 15), and small skeletal size. Basal tusk circumference of 470 mm (Zakrevska, 1935; equivalent to approx. diameter of 149 mm) is comparable to 138 mm in the female Edersleben skeleton (Garutt and Nikolskaya, 1988), and smaller than mature M. trogontherii and M. meridionalis males (see text). Male gender of Chembakchino skeleton determined from pelvic proportions (A. Rezvyi, pers. comm., 2009). b Edersleben skeleton part of ‘meridionalis-trogontherii’ transitional group, hence low dental formula (see text). c Unclear whether published plate count includes talons. d Growth incomplete (epiphyses unfused). e Skeletal shoulder height for unmounted skeletons estimated here as mean of regressions from humerus, radius and femur lengths (see below and Table 16); or estimated by Shpansky et al. (2008a,b) (S) or Bajgusheva and Titov (2007) (B). Specimens not asterisked measured directly on mounted skeletons. Living shoulder height was greater e see text. at its type locality of Süssenborn (Guenther, 1969; Lister and Joysey, various localities, such as Tiraspol, Russia, and Pakefield, England 1992), Tiraspol (Dubrovo, 1971) and elsewhere. (Nikiforova et al., 1971; Parfitt et al., 2005). The model of in situ In recent years, several important new discoveries of M. tro- European mammoth evolution would predict a series of interme- gontherii have added significantly to this record (Table 1), and diates in the transition between the two forms (e.g. Adam, 1961). allow comparison with the West Runton individual. A second However, there is accumulating evidence that M. trogontherii arose skeleton from Azov, southern Russia, has been recovered in eastern Asia as early as 2.0e1.5 Ma, based especially on the in situ (Bajgusheva and Titov, 2007). Another important skeleton, from find of trogontherii-like molars in a layer with an estimated date of Chembakchino on the Irtysh river, was described by Kosintsev ca. 1.7 Ma at Majuangou, China (Wei et al., 2003; Zhu et al., 2004). et al. (2004). Also on the Irtysh, near Pavlodar, a partial skeleton This implies that the first occurrences in Europe represent an was described by Shpansky et al. (2008a). A cranium, previously immigration event, so that a sharper biostratigraphic transition discovered very close to this site, was described as Phanagorolox- would be expected there. The possibility of temporal overlap and odon irtyshensis sp. nov. (Shpansky, 2005), but may also represent even hybridisation between the two forms in Europe has also been M. trogontherii. Finally, a skeleton with well-preserved cranium raised (Lister and Sher, 2001; Lister et al., 2005), based especially on was discovered at Kostolac, Serbia in 2009. In cranial and dental their apparent co-occurrence at Siniaya Balka, SW Russia, where morphology the specimen appears to conform to M. trogontherii, the apparently earliest European trogontherii-like form is dated to and awaits description. Skeletons here accepted as referable to M. ca. 1.0e0.9 Ma (but see Baigusheva and Titov, 2010). trogontherii are listed in Table 1, based on a combination of molar The West Runton mammoth, representing one of the earliest plate count (typically 17e22 in M3), high molar crown, and large and most complete records of M. trogontherii in western and central skeletal size (see later). The common name ‘steppe mammoth’ has Europe, is therefore of considerable significance in testing different been widely used for this species, but this term is avoided here as models of mammoth evolution. This paper describes the West its habitat evidently varied, and at West Runton the species is Runton remains, assesses them in terms of number of individuals unequivocally associated with regional temperate forest (Stuart represented, completeness, taxonomic allocation, ontogenetic age and Lister, 2010b). and sex, and discusses morphological and metric features in The latest remains of M. meridionalis in Europe, and the earliest comparison with other key specimens of the mammoth lineage. It of M. trogontherii, are generally regarded as occurring around the also briefly reassesses other elephantid remains found at West Early to Middle Pleistocene transition, in the age range ca. Runton. 0.9e0.6 Ma (Ferretti, 1999; Lister et al., 2005). Specimens of late M. meridionalis in the interval 0.9e0.8 Ma occur in the samples 2. Materials and methods from Dorst (the Netherlands) and Dorn-Dürkheim 3 (Germany) (Franzen et al., 2000; Lister et al., 2005; van Essen, in prep.). The Terminology of bones is given in Fig. 1; see Mol and van Essen species has at times been recorded from later sites, of early Brunhes (1992) for description of the osteology of Mammuthus. Bones age ca. 0.8e0.6 Ma, including West Runton, and Voigtstedt/Eder- were identified by comparing them with an Asian elephant skel- sleben (Germany) e these will be discussed below. In the same eton (University Museum of Zoology, Cambridge) for reference. interval, M. trogontherii appears in central and western Europe at Most elements could be identified without difficulty; serially Download English Version: https://daneshyari.com/en/article/7453317

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