1This report describes the left and right molar (M3) of a steppe mammoth, Mammuthus trogontherii (POHLIG, 1885), which have been discovered while digging a trench through the ancient maar of Nolhac in 1986. The molars have been extracted from the cranium, which remained in situ at the locality after it was discovered. The morphology and measurements of these molars prove that they belong to the Middle Pleistocene mammoth species Mammuthus trogontherii. Prior to this discovery, these remains have been identified by Surmely (1993, p.43), who assigned them to the southern mammoth, Mammuthus meridionalis (NESTI, 1825) without given a description. In addition, René Liabeuf collected a complete patella of most likely the same individual in 1986, not far from the site where the skull was found. In August 2008, a team of the Musée Crozatier in Le Puy‑en‑Velay excavated the remains of the cranium from which the molars were extracted at time of discovery (fig. 1). Today, this skull rests in a plaster jacket in the museum, in order to be restored in the near future.
2The village of Nolhac is located on a basaltic plateau, which is part of the geological system of the Devès (Bonifay & Liabeuf, 1994). Approximately 2 million years ago the volcano of Nolhac erupted and formed the maar of Nolhac. During this process, the crater with a diameter of 1,500 meters was filled with water and subsequently transformed into a lake. An outlet in the southwest corner (towards the village of Marminhac) caused the lake to form sediments.
3Aymard (1852) and Boule (1892) described Pleistocene remains, such as molars of horses from the maar of Nolhac.
4During the construction of a purification station in the center of the maar of Nolhac, Bonifay (1994) analyzed the site called Nolhac Briard from 1988 to 1994. Excavations there revealed lithic tools which did not correspond with the layer in which the mammoth cranium was embedded.
Fig. 1: The excavation of the cranium of the Nolhac Mammoth in the summer of 2008.
Photo: Rémi Barbe - Zoom43.fr.
5The left and right last molars (M3), which have been extracted from the cranium in 1986, are well preserved. Possessing the original number of plates or lamellae, they are almost complete. Both molars are not only large and robust, but their morphology also has all characteristics of upper molars in Mammuthus: the occlusal surface is both on the length and width axis convex. The crown is relatively width and high. The curvature of the crown, measured alongside the axis, is convex on the bucal and concave on lingual side.
6The total number of lamellae is 19, which compares perfectly to all published data on upper M3 in literature in the number of lamellae, varying from 14 to 21 in M3 of Mammuthus trogontherii (tab. 1). The enamel of the lamellae is folded and measures 2 mm in thickness, the maximum crown height and the maximum width (tab. 2 & 3) indicate low crowned molars.
7The morphology, measurements and number of plates indicate that the molars of the Nolhac mammoth should be assigned to Mammuthus trogontherii. Whereas the molars are higher crowned than in its ancestor, Mammuthus meridionalis, the enamel thickness is less, while the lamellar frequency is much higher than in Mammuthus meridionalis.
Tab. 1: Number of lamellae in the upper molars of three European mammoth species, Mammuthus meridionalis, Mammuthus trogontherii and Mammuthus primigenius, based on published data
Maglio, 1973, Mol & Van Essen, 1992
Tab. 2: M3 sup. dex. Notes and biometric data (in mm).
Tab. 3: M3 sup. sin. Notes and biometric data (in mm).
8Both molars belong to a large individual, as the diameter of the alveoli of the cranium is relatively large, holding heavy tusks with a large diameter. It is well known that there is a significant sexual dimorphism in both extinct and extant elephants, because male individuals are generally speaking much larger than females. The tusks in male individuals, especially at high age, have a large diameter and are twisted spirally, which is similar to the Nolhac specimen. Thus, we identify the Nolhac mammoth as a male individual.
9The preliminary measurements of the diameter of the alveoli gathered in the field indicate that the Nolhac mammoth was carrying large tusks. Together with the extreme measurements of the complete patella (table 4), we have the impression that the Nolhac mammoth was a large individual, measuring at least 350 cm at his highest point (shoulder height) of the backbone.
10The individual age of a mammoth can be accurately estimated based on data of the dental elements and their stage of wear gained from elephants today. The process of individual age estimation was intensively studied by Laws and published in 1966. Along with other documentation, Laws’ effort is used in paleontology in order to estimate the individual age of a mammoth species of which the molars are preserved. Comparison of mammoth molars with identical molars of the African elephant at the same stage of wear will provide an Age Group (Laws, 1966), which enables the determination of the individual age at time of death of the mammoth species, expressed in AEY (African Elephant Years).
11The left and right upper M3 and their stage of wear of Mammuthus trogontherii from the Nolhac locality can be placed in Laws (1966) Age group XXIII - XXIV given 43 ± 2 AEY to 45 ± 2 AEY. About 25 percent of the left and right molars have been used during life of the Nolhac mammoth, meaning that the Nolhac specimen died at an age of minimum 41 and maximum of 47 African Elephant Years.
12In 1885, Hans Pohlig provided the original description of the steppe mammoth without given an accurate diagnosis. Pohlig (1888-1889) considered the steppe mammoth as an intermediate species between Mammuthus meridionalis and Mammuthus primigenius. Its scientific name is based on the beaver-like animal, Trogontherium cuvieri, which was often found together with this mammoth species in the same deposits. As its popular name indicates, this mammoth species inhabited the cold and dry steppe of the Middle Pleistocene, which is confirmed by associated fauna elements, consisting of animals adapted to the steppe environment. Examples of classical steppe sites are Süssenborn in Germany and Tiraspol in Moldavia.
13The molars of the steppe mammoth are higher crowned than its ancestor, Mammuthus meridionalis, which is assumed to be an adaptation of a grazer, whereas Mammuthus meridionalis is considered a browser. Also, the length of the mandible in Mammuthus trogontherii is shorter than in Mammuthus meridionalis.
14All in all, the steppe mammoth was not only a large animal equipped with large spirally curved tusks, but also with a great sexual dimorphism in this species.
15Several mammoth skeletons have been identified as Mammuthus trogontherii. The mammoth of the late Early Pleistocene or early Middle Pleistocene of Edersleben (Spengler Museum, Sangerhausen, Germany), initially described as Mammuthus trogontherii is now considered to be a female individual of Mammuthus meridionalis. Other skeletons from the European part of Russia which have been assigned to Mammuthus trogontherii should now be considered to be the earliest representatives of the Late Pleistocene Mammuthus primigenius, the woolly mammoth.
16One of the most recent discoveries of the steppe mammoth is the more or less complete skeleton of the so-called West Runton Elephant from West Runton, (East Anglia, England) made in 1993, (Stuart, no date and Stuart 1996). The skeleton belongs to an old male individual which stood approximately 4 meters at shoulder height. It has been excavated, together with remains of an associated fauna, from fluviatile sediments dated around 700.000 – 600.000 years old.
17Like Mammuthus meridionalis the steppe mammoth could reach massive body mass proportions. Hundreds of isolated bones and molars of Mammuthus trogontherii, from the classical site Mosbach near Mainz/Wiesbaden, are stored in the collections of the Natural History Museum of Mainz, Germany. These remains identified as Mammuthus trogontherii date from the Middle Pleistocene and originate from river sediments. Some of these remains, such as a left humerus, indicate that the steppe mammoth was a giant. The maximum length of this humerus is 1450 mm and the maximum width of the distal epiphysis measures 370 mm. A complete pelvis of steppe mammoth was excavated from the ‘Mosbach-Sande’ before the Second World War had begun. Again, the measurements indicate a large (male) individual. The maximum width is 1830 mm.
Tab. 4: Patella dex., biometric data (in mm).
Plate 1: Nolhac Mammoth, Mammuthus trogontherii (POHLIG, 1885), M3 sup. dexter. Musée Crozatier, Le Puy-en-Velay, 2008-10-2-NOL. Top: Bucal view, Under: Occlusal view.
Plate 2: Nolhac Mammoth, Mammuthus trogontherii (POHLIG, 1885), M3 sup. sinister. Musée Crozatier, Le Puy-en-Velay, 2008-10-1-NOL. Top: Bucal view, Under: Occlusal view.
18Remains of the steppe mammoth are well known from several classical Middle Pleistocene sites in Europe. Particularly the molars are well known, while post‑cranial remains are rare. One of the best known specimens of the steppe mammoth is the partial skeleton of the so-called West Runton Elephant in the collections of the Norwich Castle Museum, (Norwich, England). The skull with molars and one tusk of the West Runton Elephant are well preserved and provide the best known example for comparative studies of the Nolhac specimen. These studies are planned for 2009. In France however, Mammuthus trogontherii is rare, thus the Nolhac mammoth will provide additional information on the evolution and distribution of the steppe mammoth in Europe. Moreover, the Nolhac mammoth will shed more light on the geological age of the sediments which filled up the ancient maar, where the steppe mammoth was embedded.
Plate 3: Nolhac Mammoth, Mammuthus trogontherii (POHLIG, 1885), Patella dexter. Musée Crozatier, Le Puy-en-Velay, 2008-10-3-NOL.
Top, left: cranial view, Top, right: caudal view. Middle, left: Lateral view, Middle right: axial view. Under: Ventral view.
19Comparative studies on the upper molars of the Nolhac mammoth are scheduled in Germany and England. Extensive collections with isolated molars of Mammuthus trogontherii are stored in the Research station of Quaternary Paleontology Weimar of the Senckenberg Research Institute and Natural History Museum, (Weimar, Germany). In the collection of this institute the lectotype of Mammuthus trogontherii is stored, a M3 dex. under catalogue number IQW 1965/2925 (Kahlke, 1990). The results of a comparative study on the molars of Mammuthus trogontherii will shed light on the evolutionary stage of the Nolhac specimen and this might give information on its geological age with is not clear for the time being.
20As soon as the work on the 2008 excavated cranium begins, anatomical analyses on the skull morphology of Mammuthus trogontherii will follow. This may shed a new light on the evolutionary stage of the Nolhac mammoth.
21At the Old Dominion University (Virginia, USA), a sample of the Nolhac mammoth is under study by Dr. Alex Greenwood, who is aiming to find traces of mammoth DNA. When these are found, he might be able to detect the sequence.
22The aim of the excavation campaign in 2008 was the sedimentary layers of the Nolhac maar. For palynological, sedimentary and micro-mammal purposes, all layers have been screen washed for determination. In the process, bone fragments of an unidentified large mammal were collected in a second layer beneath the mammoth skull.
23Palynological analyses will carried out, Dr. Evgenia Markova (Institute of Plantation and Animal Ecology, Yekaterinburg, Russia) will lead the micro-mammal analyses and Prof. Henri Buscail (LVEEM, Le Puy-en-Velay) is evaluating the X-ray analyses.
24The remains of the Nolhac steppe mammoth are catalogued (2008-10-1-NOL and 2008-10-2-NOL) and stored in the Musée Crozatier, (Le Puy-en-Velay, France). The originals have been molded.
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