Francien I. B r a b e r 1, Dick Mol1 & John de Vo s 2 1 N a t u u r museum Rotterd a m 2 N a t u r a l i s , L e i d e n

On remains from the Netherlands: an ove r v i ew

B r a b e r, F.I., Mol, D. & De Vos, J., 1999 - On mastodon remains from the Netherlands: an overview - in: Reumer, J.W. F. & De Vos, J. (eds.) - EL E P H A N T S H AV E A S N O R K E L! PA P E R S I N H O N O U R O F PA U L Y. SO N D A A R - DEINSEA 7: 55-65 [ISSN 0923-9308]. Published 10 December 1999

The purpose of this paper is to briefly describe two unknown molars of Anancus arv e r n e n s i s (CR O I Z E T E T JO B E RT, 1828). One specimen was trawled from the Oosterschelde and the other was found in a clay pit near the village of Maalbeek, near Tegelen, The Netherlands. Also, an overview of the known molars of A. arv e r n e n s i s and Mammut borsoni (HAY S, 1834) found in the Netherlands is given.

Correspondence: Francien Braber and Dick Mol, Natuurmuseum Rotterdam, P.O.Box 23452, NL-3001 K L Rotterdam, The Netherlands. John de Vos, Naturalis - National Museum of Natural History, P. O . Box 9517, NL-2300 RA Leiden, The Netherlands.

Keywords: A n a n c u s, M a m m u t, Plio-, Early Pleistocene, Oosterschelde, The Netherlands

I N T RO D U C T I O N Anancus arvernensis (CR O I Z E T E T JO B E RT, A n a n c u s teeth, which results in a comparable 1828) had a wide Eurasian geographical range dating of the fossils to 1,9 million years old and existed from the until the Early (Moraal 1998, Boeuf 1993). This means that Pleistocene. Its largest expanse was during the the A n a n c u s fossils found in the Oosterschelde Pliocene. The oldest known fossils of A n a n c u s belonged to the stratigraphically youngest a rv e r n e n s i s were found in Miocene deposits representatives of the species. A n a n c u s in Spain. A l a rge amount of fossils of A n a n c u s became extinct in the Early Pleistocene. was found in Bulgaria in the region of Anancus arv e r n e n s i s had a bunodont dentition Dorkovo in 1985, in a Pliocene deposit and had two tusks pointing straight forward (Thomas et al. 1986). These dental fossils are (Fig. 1). The shoulder height of the characterised by a rather primitive morpholo- was about 2 m. The mandibles were short and g y, which either indicates an Early Pliocene the molars had nipple-shaped cusps arranged form or an intraspecific variation of the spe- in transverse rows. These bunodont molars cies. Other localities where Anancus arv e r n e n - suggest a soft diet of leaves, fruit and twigs. s i s was found are in France (Chilhac), Italy The feet were made to walk on soft soil, ( Valdarno) and the Netherlands (Maalbeek, which means that A n a n c u s was an inhabitant Liessel, Schelde estuary). In France and in the of the moist woodlands. In the fossil record Schelde estuary the remains of Anancus arv e r - the crown became higher, the valleys n e n s i s were found in association with between the cusps and rows were filled with Mammuthus meridionalis. Late representatives cement and the transverse rows or lophids of A n a n c u s are characterised by molars with increased in number. The trend developed cementum in the valleys between the cusps. from soft food towards material like Both localities in France and the Netherlands grasses, requiring grinding teeth (De Vos et show this identical late evolutionary stage of a l. 1998).

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Figure 1 Reconstruction of Anancus arv e r n e n s i s.[ i l l u s t ra t i o n : Ko Sturko p ]

R E S U LT S island, this is the most probable locality. T h e N ew material of Anancus arve r n e n s i s molar became part of the private collection of Anancus arv e r n e n s i s is known from the pro- med.dr J.L. Braber in the sixties, when it was vince of Zeeland, in the south-western part of given to him as a token of appreciation for the Netherlands. Detailed descriptions and medical treatment. figures of dental remains of Anancus arv e r - n e n s i s are given by Schreuder (1944, 1945) The fossils from the Oosterschelde demon- and by Hooijer (1953, 1991). Van Essen & strate a typical state of fossilisation: when Mol (1996) and De Vos et al. (1998) gave an tapped on with a hard object they produce a overview on the Early Pleistocene - high-pitched sound. The molar in the Braber ian remains from the Oosterschelde in the collection also has those characteristics and is Zeeland province. Most of the known very well preserved (Table 1, Fig. 2). Only at remains of Anancus arv e r n e n s i s are kept in the right front side a cusp has been broken. Naturalis, the National Museum of Natural The colour of the fossil is black for the ena- History in Leiden (see Appendix). One molar mel and dark brown for the cementum. of Anancus arv e r n e n s i s belongs to the Braber Between the cusps there are relatively heavy collection (Delft, The Netherlands) and bears cementum deposits. The molar is slightly the number B0100. This molar was unpublish- worn at the conules of the protocone and at ed so far. The exact locality where this the conelets of the paracone, the protocone A n a n c u s molar was found is not known, but itself is missing (broken). The other cones, it was dredged from the bottom of the sea by conelets and conules are unworn. The roots a vessel from a fisherman from the island of of the tooth are missing. The molar is an M3 Schouwen Duiveland. Bryozoa are clearly dex. with 6 lophs of two cones, of which the recognisable on the surface of the molar. 6t h loph is much smaller than the other five. Since there are other finds of A n a n c u s k n o w n In the first loph both cones are accompanied from the Oosterschelde which borders the by two conelets (buccal side of the sulcus) or

56 BRABER et al.: from the Netherlands

two conules (lingual side of the sulcus). In Tegelen, much older such as the 2n d , 3r d and 4t h loph there is only one cone- Ta p i rus arv e r n e n s i s and Anancus arv e r n e n s i s let or conule. In the two last lophs each cone are known (We s t e r h o ff et al. 1998). Only one has two conelets and two conules, except for set of Ta p i r u s molars (Kortenbout van der the smallest cone, which has only one cone- Sluijs 1960) and one molar of A n a n c u s let. The cones and conules do not form a (ST96438) were found. Often we come straight ridge, but do alternate each other in a across both species mentioned in the faunal way that the lingual cone is situated more to list of Tegelen. There has been a lot of dis- the front than the buccal cone of the same cussion on the age of the Maalbeek Pit l o p h . ( We s t e r h o ff et al. 1998). According to many researchers the clay of the Maalbeek Pit is The Anancus arve r n e n s i s molar fro m placed in the Eburonian (e.g. Zagwijn 1963). M a a l b e e k A famous site of Early Pleistocene faunal The molar of A n a n c u s was collected by the remains is Tegelen in the province of museum staff of the Leiden Geological L i m b u rg, The Netherlands. Several clay pits Museum on February 23, 1960. near Tegelen have yielded large quantities of G. Kortenbout van der Sluijs, under the fossils. Most of them originate from the same leadership of professor I. M. van der Vlerk, at deposits (Kortenbout van der Sluijs & that time was doing a research project on the Zagwijn 1962). The main fauna of Tegelen is Tegelen fauna. Kortenbout van der Sluijs known from the stage Tiglian C 5, often writ- wrote on that day in his Tegelen diary (which ten as TC5, and dated with an age of approxi- is now in the collection of Dick Mol, Fig. 3): mately 1.6 million years. From a site called the Van Cleef Pit or Maalbeek Pit near the ‘Met Prof.v.d. Vlerk eerst naar Russel- village of Maalbeek in the neighbourhood of Tiglia. We krijgen hier een aantal beende- ren, o.a. een stuk gewei van E u c l a d o c e ro s Ta ble 1 Measurements in cm of the Anancus arv e r n e n s i s en een snijtand van een paard. Smeets is molar B0100 (coll. B ra b e r ) . The enamel thickness, measured at echter ziek, zodat een deel van de vondsten the side where the protocone has been broke n , v a ri e s ontvindbaar is. Vervolgens naar Maalbeek. b e t ween 0.53 cm and 0.81 cm. Hier krijgen we een aantal kleinere botjes, o.a. stukjes kies van C e rvus rh e n a n u s en een kies van Anancus arv e r n e n s i s. Is de klei in deze groeve toch Onder- Ti g l i e n ? ’

Translation: ‘ With Professor van der Vlerk first to R u s s e l - Tiglia. Here we obtain several bones, among which part of an antler of E u c l a d o c e ro s and an incisor of a horse. Smeets, however, is ill, so a part of the finds is untraceable. After that to Maalbeek. Here we get some smaller bones, among which dental remains of C e rvus rh e n a n u s and a molar of Anancus arv e r n e n s i s. Is the clay in this pit Lower Tiglian after all?’

Westerhoff et al. (1998) wrote in their extensi- ve paper on the Maalbeek Pit that the molar was found in situ (1998: 61). This is not quite

57 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

a

b

Figure 2 The upper M3 dex B0100 of Anancus arv e rn e n s i s; a side view and b occlusal view (anterior to the left). [ p h o t o : Esmee van Bokhove n ]

58 BRABER et al.: mastodons from the Netherlands

consistent with the notes Kortenbout van der resulted in alternative ideas about the Sluijs wrote in his diary (see above) on his stratigraphical age of the fauna from project on the Tegelen fauna. The workers in the Maalbeek. The clay with the mastodont pit donated the molar to him. We s t e r h o ff et al. molar is here referred to the Tiglian B (1998) gave some interesting information (p. 62): pollen zone. The remains might be ‘The new exposures in 1993 and 1994, slightly older and can probably be assigned additional palynological research and a to the Tiglian A pollen zone. The new reappraisal of the primary data have results indicate that the mammal fauna

Figure 3 The section in the diary of G. K o r tenbout van der Sluijs written on Febru a r y 23, 1960 (Collection Dick Mol).

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mentioned above is not younger but older Ta ble 2 Measurements in cm of the Anancus arv e rn e n s i s than the fauna from the Tegelen pits.’ molar ST96438 from the Maalbeek Pit (Coll. N a t u ra l i s , National Museum of Natural History, L e i d e n ) . The enamel This means that Kortenbout van der Sluijs, t h i c k n e s s , measured on the occlusal of the molar, v a ri e s when he wrote his notes in his Tegelen diary, b e t ween 0.29 cm and 0.65 cm. was correct in his assumption about the dating of a Lower Tiglian age of the Maalbeek Pit. The molar (ST96438) now belongs to the collection of Naturalis in Leiden. It is the only A n a n c u s fossil ever found in Maalbeek. This is the first time it is described, measured and depicted (Table 2, Fig. 4). ST96438 is an upper M1 dex., which is very worn down. On the lingual side of the molar all cusps are totally worn. On the buc- cal side five lophs can be distinguished, all worn.

An ove r v i ew of the Anancus arve r n e n s i s molars found in the Netherlands The Appendix provides an overview of all the molars and molar fragments known from the Netherlands. Until now there were found ten (almost) complete molars and 26 molar frag- ments of Anancus arv e r n e n s i s. They all show comparatively heavy cementum deposits in the collection of the Natuurmuseum between the cusps and therefore belong to the Rotterdam, is of unknown provenance. Molar late evolutionary stage. The localities were Li 1001 was found in 1998 at a depth of 20 the A n a n c u s and the M a m m u t molars were m e t e r. It is a crown of a left m3 with five found in the Netherlands and in the North ridges, of which the first four are called Sea are indicated on the map (Fig. 5). lophids and the last the talonid. It is heavily worn, especially at the buccal side, which An ove r v i ew of the M a m mut bors o n i indicates that it belonged to an old animal. molars found in the Netherlands The second molar from Liessel was found in Mammut borsoni ( H AY S , 1834) is a zygodont 1990 at a depth of 22 meters. It is the com- mastodon known from the Pliocene of Europe plete crown and a part of the root of a left m3 and Northern Asia. The molars have sharp (Peters et al. 1991). transversal ridges, derived from two fused cones, specialised for cutting the food like The stratigraphy of the Liessel site is very scissors. There are only two or three molars complicated, because the sediment was of Mammut borsoni found in the Netherlands. dredged from the pit. The fossils derived Two were found at the locality Liessel, a from the pit are diverse; remains of marine gravel pit in the province of North Brabant; mammals dated into the Miocene and fossils Li 1001 is in the collection of the Museum of terrestrial mammals of Pliocene age. A l s o Jan Vriends in Asten and the other one in the plant remains from the Pliocene were reco- collection of mr. Nies in Deurne. They were vered. A Pliocene age for the molars of found at about 100 meters distance from each Mammut borsoni is most convenient, like the o t h e r. The third molar (NMR 698), which is other terrestrial mammals and the plant seeds.

60 BRABER et al.: mastodons from the Netherlands

a

b

c

Figure 4 The upper M1 dex (ST96438) of Anancus arv e rn e n s i s from Maalbeek (The Netherl a n d s ) ; a side view, b occlusal view and c a drawing of the surface of the undamaged enamel (anterior to the ri g h t ) .

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Figure 5 Sites (underlined names) of all the known molars of Anancus arv e r n e n s i s and Mammut bors o n i in the Netherl a n d s . [ i l l u s t ra t i o n : Jaap van Leeuwen]

The third specimen (NMR 698) is only a part Mammut borsoni is often found associated (two ridges) of a lower m3. There is no loca- with Anancus arv e r n e n s i s. They must have lity information available. Although Vo n had a different diet according to their denti- Koenigswald (1950) interpreted it as a fossil tion; Mammut borsoni cutting branches and found in the Oosterschelde, this is improba- twigs with its zygodont teeth, Anancus arv e r - ble. There are no traces of sea-life on the sur- n e n s i s with its bunodont teeth browsing on face. The fossils found in the sea always have leaves, both in the moist woodlands at the barnacles or bryozoans stuck to the surface. river banks. Another problem is that Mammut borsoni i s known from the Pliocene and the finds of the Oosterschelde are younger, at least of Early Pleistocene age.

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C O N C L U S I O N S The only A n a n c u s molar which is known Anancus arv e r n e n s i s and Jaap van Leeuwen from Maalbeek is an upper M1 dex. Its geo- for drawing the map of the Netherlands. logical age is Tiglian B. Kortenbout van der Sluijs (1960) was the first author to recognise R E F E R E N C E S that the Maalbeek fauna was older than the Boeuf, O., 1993 - Il était une fois… Il y a près de main fauna of the famous Tegelen clay pit. 2.000.000 d’années à Chilhac, Haute-Loire, France - All the published molars and molar fragments Amis de Chilhac (Association) of Anancus arv e r n e n s i s known from the De Vos, J., Mol, D. & Reumer, J.W. F., 1998 - Early Netherlands are listed in this paper and all the Pleistocene mammalian remains from the molars show the heavy cementum deposits Oosterschelde or Eastern Scheldt (province of between the valleys of the cusps. This indica- Zeeland, The Netherlands) - Mededelingen tes that they all had the same evolutionary Nederlands Instituut voor Toegepaste stage and therefore we can conclude that Geowetenschappen TNO 60: 173-185 Anancus arv e r n e n s i s became extinct in the H o o i j e r, D.A., 1953 - On dredged specimens of A n a n c u s , Early Pleistocene in North-western Europe, A rc h i d i s k o d o n and E q u u s from the Schelde Estuary, probably during the Tiglian B. Mammut The Netherlands - Leidsche Geologische b o r s o n i is often found associated with Mededelingen 19: 255-256 Anancus arv e r n e n s i s, but only restricted to H o o i j e r, D.A., 1991 - Fossielen vissen voor de weten- the Pliocene, while A n a n c u s is also known schap; veertig jaar Kor en Bot (1951-1991) - from the Early Pleistocene (Mol & Van Essen Kroniek van het Land van de Zeemeermin 1996). In Dorkovo (Thomas et al. 1986) the (Schouwen Duiveland): 1-16 two proboscideans were also found associa- Kortenbout van der Sluijs, G. & Zagwijn, W.H., 1962 - ted, they are of Pliocene age and the larg e An introduction to the stratigraphy and geology of the molars of Anancus arv e r n e n s i s represent the Tegelen clay-pits - Mededelingen Geologische more primitive stage. In Liessel, where the Stichting NS 15: 31-37 stratigraphy is rather complicated and sedi- Kortenbout van der Sluijs, G., 1960 - The fossil tapir of ments could be mixed, M a m m u t and A n a n c u s Maalbeek, Netherlands - Publicaties Natuurhistorisch are both present, but it is not clear that they Genootschap in Limburg 12: 12-18 derive from the same deposit. Because Mol, D.J. & van Essen, J.A., 1990 - Mammut borsoni A n a n c u s represents the late evolutionary from the Netherlands - Lutra 33 (2): 183-186 stage in dentition, characteristic for the Early Moraal, J.M., 1998 - Genootschapsverzamelingen, de Pleistocene age, and the age of M a m m u t mastodont van A u v e rgne - Zeeland, Tijdschrift van fossils is restricted to the Pliocene in Europe, het Koninklijk Zeeuws Genootschap der We t e n - it is assumed that the layers of Late Pliocene schappen 7 (2): 69-74 and Early Pleistocene age in Liessel are Peters, A., Lammers, Th. & Mol, D., 1991 - mixed up and that the remains of the two Mastodonten-kiezen uit Liessel (Noord-Brabant) - Proboscideans derived from different layers. Cranium 8 (2): 89-96 S c h r e u d e r, A ., 1944 - Upper-Pliocene out of AC K N OW L E D G E M E N T S the Scheldt and the Lower Rhine - Leidsche The authors would like to express their Geologische Mededelingen 14: 40-58 gratitude and admiration to dr Paul Sondaar S c h r e u d e r, A ., 1945 - De mastodont en de olifant onder for his continuous and ongoing enthusiasm to de “zwarte fossielen” uit de Zeeuwsche wateren - develop the Pleistocene vertebrate palaeonto- Verhandelingen van het Geologisch en logy both on a national and global scale with Mijnbouwkundig Genootschap 14: 437-448 great skill. We would like to thank dr Jelle Thomas, H., Spassov, N., Kodjumdgieva, E., Poidevin, Reumer for his valuable comments on this J.L., Popov, V., Sen, S., Ta s s y, P. & Visset, D., 1986 - p a p e r, Ko Sturkop for the reconstruction of Résultats préliminaires de la première mission

63 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

paléontologique franco-bulgare à Dorkovo (arrondis- Nederland - Geologie en Mijnbouw 12 (1): 14-15 sement de Pazardjik) Bulgarie - Comptes rendus de We s t e r h o ff, W.E., Cleveringa, P., Meijer, T., Van l’Académie des Sciences Paris 302, II (16): 1037- Kolfschoten, M. & Zagwijn, W.A., 1998 - The Lower 1 0 4 2 Pleistocene fluvial (clay) deposits in the Maalbeek pit Van Essen, H. & Mol, D., 1996 - Plio-Pleistocene near Tegelen, The Netherlands - Mededelingen proboscideans from the southern bight of the North Nederlands Instituut voor Toegepaste Sea and the Oosterschelde, The Netherlands - in: Geowetenschappen TNO 60: 35-65 Shoshani, J. & Ta s s y, P. (eds.) - The Proboscidea, Zagwijn, W.H., 1963 - Pollen-analytic investigations in Evolution and Palaeoecology of Elephants and their the Tiglian of the Netherlands - Mededelingen van de relatives - Oxford University Press: 214-224 Geologische Stichting NS 16: 49-69 Von Koenigswald, G.H.R., 1950 - Voorlopige medede- ling omtrent het voorkomen van Mastodon borsoni in r e c e i ved 12 July 1999

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APPENDIX An ove r view of the Anancus arv e r n e n s i s m o l a r s and molar fragments found in the Netherl a n d s .

65 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

DEINSEA - ANNUAL OF THE NATURAL HISTORY MUSEUM ROTTERDAM P. O . B o x 2 3 4 5 2 , N L - 3 0 0 1 K L R o t t e r d a m T h e N e t h e r l a n d s

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