Dick Mol1, G e rt D. van den Bergh2 & John de Vo s 2 1N a t u u r museum Rotterd a m 2N a t u r a l i s , L e i d e n

Fossil proboscideans from The , the and the Oosterschelde Estuary

Mol, D., van den Bergh, G.D. & de Vos, J., 1999 - proboscideans from The Netherlands, the North Sea and the Oosterschelde Estuary - in: Haynes, G., Klimowicz, J. & Reumer, J.W. F. (eds.) – MA M M O T H S A N D T H E MA M M O T H FA U N A : ST U D I E S O F A N EX T I N C T EC O S Y S T E M – DEINSEA 6: 11 9 - 1 4 6 [ISSN 0923-9308]. Published 17 May 1999.

This paper reviews the most important findings of proboscidean from The Netherlands, both f r o m the mainland as well as those dredged by fisherman from the estuaries and the Southern Bight of the North Sea. The oldest, , proboscidean remains from The Netherlands are a few isolated molars of Mammut borsoni from Liessel. Also Anancus arv e r n e n s is occurs at this locality. An Early fauna with A. arv e r n e n s i s and M. meridionalis has been dredged from the Oosterschelde E s t u a r y. The Oosterschelde fauna, with an estimated age of 1.9 myr, is slightly older than the vertebra- te fauna from the clay pits at Tegelen. The postcranial fossils of A n a n c u s and Mammuthus meridiona - l i s , both heavily mineralized, can be easily distinguished based on size differences, but also on morphological grounds. Whereas A n a n c u s has its latest occurrence in the Oosterschelde assemblage, M. meridionalis continues well into the Middle Pleistocene and is known from Dutch mainland locali- ties as well as from the North Sea. From the North Sea also a few molars of M. tro g o n t h e r i i have been dredged. Fossils of the straight-tusked , E. antiquus, are rare. Some of the earlier remains of this species occur in mainland sites and have a late Middle Pleistocene age. Late Pleistocene probosci- deans are restricted to M. primigenius and E. antiquus, both showing a light degree of fossilization. While the woolly is known from a wealth of shallow localities in The Netherlands, it is especially common amongst fossils dredged from the North Sea. Late Pleistocene E. antiquus remains are rare.

Fossiele olifantachtigen uit Nederland, de Noordzee en het Oosterschelde bekken – In dit artikel wor- den de belangrijkste vondsten van olifantachtigen uit Nederland beschreven, zowel van het vasteland als het opgeviste materiaal uit de zeearmen en de Noordzee. Het oudste, Pliocene olifantmateriaal zijn enkele losse molaren van Mammut borsoni uit Liessel. Uit deze vindplaats is ook Anancus arv e r n e n s i s bekend. Een vroeg-Pleistocene fauna met A. arv e r n e n s i s en M. meridionalis is opgevist uit de Oosterschelde. De Oosterschelde fauna is met een leeftijd van ca. 1,9 miljoen jaar iets ouder dan de fauna van Tegelen. Het postcraniale materiaal van A n a n c u s en Mammuthus meridionalis, beide zwaar gemineraliseerd, kan worden onderscheiden op basis van maatverschillen en morfologie. Te r w i j l A n a n c u s in de Oosterschelde zijn jongste voorkomen heeft, komt M. meridionalis voor tot in het Midden Pleistoceen, zowel op het vasteland als in de Noordzee. Uit de Noordzee is ook een aantal kie- zen van M. tro g o n t h e r i i bekend. Fossielen van de bosolifant, E. antiquus, zijn zeldzaam. Enkele vroe- ge vondsten van deze soort komen van het vasteland en dateren uit het late Midden Pleistoceen. Laat Pleistocene olifanten bepreken zich tot M. primigenius en E. antiquus, beide slechts licht gefossiliseerd. Terwijl de wolharige mammoet bekend is uit veel ondiepe vindplaatsen in Nederland, is deze soort bij- zonder veelvuldig aangetroffen tussen fossielen die uit de Noordzee zijn opgevist. Laat Pleistocene overblijfselen van E. antiquus zijn zeldzaam.

Correspondence: Dick Mol, Natuurmuseum Rotterdam, P.O.Box 23452, NL-3001 KL Rotterdam, T h e Netherlands (or private: Gudumholm 41, NL-2133 HG Hoofddorp, The Netherlands); Gert D. van den B e rgh and John de Vos, Naturalis, National Museum of Natural History, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands.

Keywords: , The Netherlands, , morphometrics

119 AND THE MAMMOTH FAUNA DEINSEA 6, 1999

I N T RO D U C T I O N Fossil teeth and bones of Quaternary mam- A well-documented terrestrial fauna of ca. 1.7 mals are commonly preserved in the unconso- My old has become known from this type lidated sediments underlying the Dutch delta. locality of the Tiglian stage. The fossils can usually not be found at the exposed landsurface, but are dredged up as a Proboscidean remains constitute an important result of sand- and gravel dredging operations element from these various sources, ranging in the alluvial plains of the rivers Rhine, in age from Pliocene to Late Pleistocene. T h e Waal, and IJssel. In the North Sea majority of the fossils originate from Late h o w e v e r, fossil-bearing strata crop out on the Pleistocene layers. Though the latter are usu- seabottom at several places, and an important ally not mineralized, they are often well pre- source of fossil terrestrial in T h e served, especially those dredged from the Netherlands are fishing boats. The trawlers North Sea bottom. Mammuthus primigenius use beamtrawls that are towed over the sea is one of the most common elements in the bottom. During these operations large debris, Late Pleistocene faunas. Less well known including mammalian fossils, is collected ins- from The Netherlands are the rarer occurren- ide the nets. The trawlers that operate in the ces of Pliocene and Early Pleistocene probos- southern bight of the North Sea between cideans, such as Anancus arv e r n e n s i s, and England and The Netherlands and also in the Mammut borsoni. In addition, various collec- Schelde Estuary, have over the years brought tions comprise a fair amount of fossils of in an enormous amount of mammalian fossils Mammuthus meridionalis and anti - in this way. Because the mesh diameter of the q u u s , whereas Mammuthus tro g o n t h e r i i nets is about 5 cm, mostly larger fossils are remains are very rare in The Netherlands. brought in by the fishing boats. Many well- The scientific significance of the dredged fos- preserved dredged mammalian fossils have sils is often considered as less important, ended up in private collections, several of because the exact stratigraphic level of origin which are well documented. Also, a conside- is not known. On the other hand, the fossils rable amount of this dredged material is sto- often have an excellent state of preservation, red in Naturalis, the National Museum of and also the large amount of fossil specimens Natural History (NNM) at Leiden (formerly makes them an important source of informa- the National Museum of Geology and tion. Besides, the mapping of the North Sea Mineralogy), which institution contains one Quaternary (Cameron et al. 1984) in combi- of the largest collections in the world of fos- nation with information on the dredging loca- sils of the , Mammuthus pri - tions has made it possible to reconstruct the m i g e n i u s. Also, dredged fossils are being sold age of many fossil findings from the North in many countries, and The Netherlands have Sea (Van Kolfschoten & Laban 1995). A l s o become the top country in the world in the the subsurface geology of the Dutch delta has export of mammoth teeth. During the last 45 become increasingly well known, so that fos- years the museum in Leiden has also been sil specimens from sand and gravel dredging actively involved in the search for fossil operations can now often be placed within remains from the sea bottom, by employing a their stratigraphical context with reasonable fishing boat once every year to bring up fos- c e r t a i n t y. Also the fossil assemblages them- sils from the Oosterschelde. Finally, many selves from certain locations provide clues important fossil remains from the North Sea concerning the age of these faunas. The Early can be found in a wide scala of smaller natu- and Middle Pleistocene fossils can be distin- ral history musea throughout The Netherlands. guished from Late Pleistocene fossils because A third source of mammalian fossils in T h e of their state of preservation. It is not so Netherlands are the clay pits near Tegelen. much the color of the fossils, which is of

120 MOL et al.: Netherlands proboscideans

importance to make this distinction (Drees the dredged material from the Oosterschelde, 1986), but the degree of mineralization: the from where so far no M. borsoni material has fossils originating from older formations are been reported with certainty. This suggests heavily mineralized and have a relatively that the Liessel locality has yielded the oldest high density. They produce a high-pitched proboscidean remains in The Netherlands, sound when tapped on with a hard object. with a Pliocene age. The Late Pleistocene bones on the other hand, such as the remains of M. primigenius, Anancus arve r n e n s i s have a relatively low density and produce a This taxon has its oldest occurrence in dull sound. various Late (Late Turolian) locali- ties in Spain and had a wide Eurasian distri- In this paper we will give a brief overview of bution during the Pliocene, but became the various proboscidean remains from T h e extinct during the Early Pleistocene. T h e Netherlands. The geological context of the various A n a n c u s remains from T h e various Plio-Pleistocene fossil-bearing depo- Netherlands are thought to represent the latest sits in the North Sea and a description of the populations before this species became proboscidean remains from the North extinct. Most material originates from the Sea have already been dealt with in other Oosterschelde (Schreuder 1944, 1945, papers (Van Kolfschoten & Laban 1995, Va n Hooijer 1953), where A n a n c u s seems to be Essen & Mol 1996). Here we will give a associated with remains of Mammuthus meri - review of the proboscidean remains from T h e dionalis and other terrestrial mammals, broadly Netherlands, including the North Sea and indicative of an Early to Middle Pleistocene Oosterschelde estuary. Postcranials of age. The fossils originate from part of the Anancus arv e r n e n s i s and Mammuthus meri - Oosterschelde where the Tegelen Formation d i o n a l i s from the North Sea and Ooster- crops out (Drees 1986). However, from the schelde have not yet been described before beginning it was realised that the fauna from and they will be compared and described in the Oosterschelde was older than the famous this paper. Also attention will be put on the fauna from the clay pits near Te g e l e n accompanying faunas in which the various (Province of ). The Oosterschelde proboscideans occur. fauna has been indicated as Pre-Ti g l i a n (Hooijer 1950, 1953, 1957, Van der V l e r k 1951, Van der Feen 1968, Dumon Tak 1973). P ROBOSCIDEANS FROM T H E Because the pollen spectra from the Te g e l e n N E T H E R L A N D S clays have shown that the sequence repre- M a m mut bors o n i sents a longer period with a succession of The rarest proboscidean from T h e colder and warmer phases (Zagwijn 1963), it Netherlands, Mammut borsoni, was first is better to speak of the Tiglian Complex. T h e reported by von Koenigswald in 1950, who Tegelen vertebrate fauna is now placed in the described a lower M3 of unknown provenan- TC5 zone of the Tiglian (Van Kolfschoten & ce. The specimen is nowadays in the collec- Van der Meulen 1986), which represents a tion of the Natuurmuseum Rotterdam. More warm period and has an age of approximately r e c e n t l y, two lower M3 were discovered at 1.7 My. A n a n c u s is often included in this Liessel (Mol & Van Essen 1990, Peters et al. fauna. However, the one molar fragment attri- 1991). From this locality also Anancus arv e r - buted to A n a n c u s was not found at the same n e n s i s has been reported. Whereas locality as most of the other vertebrate M. borsoni disappears from during remains, but in the clay pit Van Cleef near the Pliocene, A. arv e r n e n s i s continues into Maalbeek, 3 km south of Tegelen (Van Essen the Pleistocene. The latter is present amongst & Mol 1996). The associated pollen spectrum

121 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

of the A n a n c u s molar indicates a cold period, Sluijs 1985) according to Zagwijn (1960, 1963) represen- - S t e p h a n o rhinus cf. etru s c u s (Mol & De Vos ting the Eburonian, which followed the 1 9 9 5 a ) warm-temperate Tiglian stage during which - E u c l a d o c e ros ctenoides, the large cervid of the famous Tegelen fauna lived. However, Tegelen (De Vos et al. 1995) based on recent studies of the clay pit - C e rvus rh e n a n u s, the middle sized deer of Maalbeek, Westerhof (personal communica- Tegelen (De Vos et al. 1995) tion to van Kolfschoten & Laban, 1995) con- - a large boar (Mol & De Vos 1996) cluded that the A n a n c u s molar predates the famous Tegelen fauna. This suggests that the There is general agreement nowadays that the Oosterschelde fauna is older than the T C 5 l a rge cervid of Senèze and Chilhac, referred stage and there is good additional evidence to as E u c l a d o c e ros senezensis, is the same for this. The Oosterschelde fauna shows great species as the large cervid from Tegelen, usu- resemblance with the fauna from Chilhac ally referred to as E u c l a d o c e ros tegulensis (Haute-Loire, ; Mol & De Vos 1996). (Germonpré 1983, Azzaroli et al. 1988, The Chilhac fauna has been dated at 1.9 My Azzaroli & Mazza 1992, Spaan 1992). In a (Bout 1979) and contains the following taxa review of the Early Pleistocene cervids from according to Boeuf (1983, 1993): Europe (De Vos et al. 1995) it was concluded - Ursus etru s c u s, the Etruscan bear that the name E u c l a d o c e ros ctenoides (NE S T I , - M e g a n t e reon megantere o n , a sabre-toothed 1841) has priority and should be used for this c a t l a rge cervid. The antler of the middle-sized - P a c h y c ro c u t a ( = H y a e n a) p e rr i e r i , the deer from the Oosterschelde consists of a hyaena of Perrier beam, one brow tine and at the end one side - N y c t e reutes megamastoides, a fox-like tine, similar to the middle-sized deer of a n i m a l Tegelen, which is known under the name - Anancus arvernensis chilhiacensis, a C e rvus rh e n a n u s DU B O I S , 1904. Spaan (1992) m a s t o d o n t made a detailed study of the Tegelen deer and - Mammuthus meridionalis, the southern concluded that C e rvus philisi SC H A U B , 1941 m a m m o t h and C e rvus pero l e n s i s (AZ Z A R O L I, 1952), see - Equus stenonis guthi, a large horse Bout & Azzaroli (1952) are synonyms of - D i c e ro rh i n u s ( = S t e p h a n o rh i n u s) e t r u s c u s, C e rvus rh e n a n u s from Tegelen. As follows the Etruscan rhino from the species lists given above, the faunas - E u c l a d o c e ros senezensis, a large deer from Chilhac and the Oosterschelde show - C e rvus philisi, a middle-sized deer great resemblance to each other, and an age - C ro i z e t o c e ros ramosus, a small deer of 1.9 myr for the Oosterschelde fauna, - Gazellospira tort i c o r n i s, a gazelle somewhat older than the Tegelen Fauna, which lacks Anancus arv e r n e n s i s, seems well The Oosterschelde assemblage contains the f u n d e d . following taxa according to the review given by Mol & De Vos (1996): In addition to the Oosterschelde, A n a n c u s - a sabre-toothed cat (Hooijer 1962, 1991) material has also been dredged from the - cf. Hyaena perrieri (Mol & De Vos, 1995a,b) Thornton Bank approximately 12 nautic miles - Mammuthus meridionalis (Hooijer 1953, o ff the coast of the province of Zeeland (51o 1962, 1981, Van Essen & Mol 1996) 35' 49" N, 03o 01'39" E). Also here A n a n c u s - Anancus arv e r n e n s i s (e.g. Hooijer 1953, occurs in association with M. meridionalis 1 9 9 1 ) and the close proximity to the Oosterschelde - a large, heavily built horse (see among suggests that they originate from the same others: Hooijer 1953; Kortenbout van der Tegelen Formation. Two A n a n c u s molar frag-

122 MOL et al.: Netherlands proboscideans

Figure 1 Po s t e r ior fragment of a left M1 of Anancus arv e rn e n s i s ( C o l l . NNM no. S t - 4 0 1 3 2 6 ) . This specimen was dredged up from the Oosterschelde Estuary near W i s s e ke rke, P r ovince of Zeeland,The Netherl a n d s . A lingual view, B occlusal view. ments are known from the Thornton Bank the same locality, the larger part fragmenta- (Coll. Mol No. 1752 and one specimen in r y, except for bones of the manus and pes. the Collection Jager at Goes). The specimen Most of this material is stored in the collec- in the Mol collection is a lower left M2 frag- tions of the NNM. Some of the carpal and ment, it was figured in Van Essen & Mol tarsal bones, typical for A n a n c u s , will be (1996: fig. 20.3). The A n a n c u s m a t e r i a l described and compared with M. meridiona - dredged from the Oosterschelde consists of l i s and M. primigenius in the last section of 39 nicely preserved molars (Fig. 1), inclu- this paper. ding elements of the milk dentition (Fig. 2), and tusk fragments. In addition, about a hundred postcranial elements originate from

123 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

Figure 2 Po s t e r ior fragment of a right DP4 of Anancus arv e rn e n s i s ( C o l l . NNM no. S t - 1 1 8 9 3 4 ) , dredged up from the O o s t e rsc helde Estuary near the Flauwe r s p o l d e r, P r ovince of Zeeland,The Netherl a n d s . A lingual view, B occlusal view.

M a m muthus meridionalis a n d was the first one who described in detail den- M a m muthus trog o n t h e r i i tal elements of Mammuthus meridionalis Mammuthus meridionalis was much larg e r from the North Sea. Based on molar measure- than Anancus arv e r n e n s i s, with which it co- ments, Van Essen & Mol (1996) intended to occurs in the Oosterschelde fauna (De Vo s e t ascribe the North Sea and Oosterschelde a l. 1996). Besides from the Oosterschelde material to one or more of the evolutionary (Fig. 3), remains of M. meridionalis h a v e stages of M. meridionalis ( Van Essen & Mol also been found in the North Sea, mainly 1996). These stages, recognised by Maglio from the Deep Water Channel. Hooijer (1984) (1973) and other authors before, have been

124 MOL et al.: Netherlands proboscideans

Figure 3 Po s t e r ior fragment of a right m3 of Mammuthus meri d i o n a l i s ( C o l l . NNM no. S t - 1 7 0 0 6 4 ) , dredged up from the O o s t e r schelde Estuary. A lingual view, B occlusal view. labelled ‘Laiatico stage’, ‘Montevarchi stage’ with the material from Chilhac, which has an and ‘Bacton stage’ (‘primitive’, ‘typical’ a n d age of approximately 1.9 myr (Bout 1970). ‘ a d v a n c e d ’ stage respectively). These succes- Three of the Chilhac skulls have M3 with 13 sive stages are characterised by increasing lamellae and one has 12 lamellae. A c c o r d i n g plate numbers, hypsodonty index and lamel- to Maglio (1973) the Laiatico stage is charac- lar frequency, and decreasing enamel thick- terised by 11 or 12 lamellae in the upper M3. ness as defined by Maglio (1973). The upper Thus, the North Sea material tends to be M3's from the North Sea have 13 lamellae slightly more advanced than the Laiatico ( Van Essen & Mol 1996). This corresponds s t a g e .

125 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

From the Deep Water Channel, a deep gully Pleistocene based on the evolutionary stage in the Southern Bight of the North Sea, (‘Bacton Stage’) of this m3. Early to early Middle Pleistocene terrestrial remains have been trawled. T h e In a paper reviewing the Pleistocene only proboscideans known from this site are: Proboscideans originating from the Early M. meridionalis and M. tro g o n t h e r i i ( H o o i j e r Pleistocene deposits from The Netherlands 1984, Mol & Van Essen 1992). A. arv e r n e n - the next three onshore sites should be men- s i s is not known from this site. In the tioned. First of all the famous Tegelen site, Oosterschelde and the Thornton Bank M . with an age of 1.7 My, has yielded in total m e r i d i o n a l is and A. arv e r n e n s i s occur in the 11 fragmentary molar remains of M. meri - same fauna (Mol & De Vos 1996). As arg u e d d i o n a l i s , which have been thoroughly stu- above, the Oosterschelde Fauna is probably died by Guenther (1986). Rutten (1909) equivalent to the 1.9 myr old Chilhac Fauna. figured and described in detail a set of Based on the absence of A. arv e r n e n s i s molars of a single individual (m1 sin. and amongst the North Sea material it is conside- dex. and M1 sin. and dex.) of M. meridiona - red likely that this material is slightly youn- l i s , which were found in 1842 in a clay pit ger than the Oosterschelde/Thornton Bank near Oosterhout (Province of Noord- material. This assumption is strengthened by Brabant). These molars were found together the fact that amongst the North Sea material with, according to Rutten (1909) remains of there are various molar fragments referable M2's and some postcranial material, proba- to Mammuthus tro g o n t h e r i i ( Van Essen & bly belonging to the same individual. Mol 1996), which species is believed to have Unfortunately it is only known where the gradually evolved from M. meridionalis. four M/m1's are kept, which is the Brabants Museum at 's-Hertogenbosch (Bois-le-Duc). In addition to molars there are numerous According to Rutten (1909), the finds from heavily mineralized postcranial remains from Oosterhout were collected in situ and have the North Sea and the Oosterschelde that an Early Pleistocene age. However, their must belong to M. meridionalis. Below we exact stratigraphical position is unknown. will describe some carpals and tarsals from Mammuthus meridionalis molars were col- the North Sea, Thornton Bank and the lected in a clay-pit called ‘Surae’, situated Oosterschelde. These can be easily distin- east of the village of Dorst (Province of guished from homologues of A. arv e r n e n s i s Noord-Brabant). The Dorst-Surae material from the Oosterschelde and Thornton Bank, was described by Van Kolfschoten (1990), based on morphological characters and size and its geological age was discussed. T h e measurements (the possibility remains that clay layer from which the fossils originated some of the early Middle Pleistocene post- was deposited during the end phase of the cranial remains actually belong to M. tro g - Leerdam Interstadial and the Dorst Stadial o n t h e r i i instead of M. meridionalis). From (the last interstadial and stadial of the M. primigenius they are distinguished by Bavelian; Zagwijn & De Jong 1984). T h e y their usually larger size and consistently would have an age slightly younger than the heavier mineralization. A well preserved Jaramillo Subzone (0.97-0.90 My), the latter complete but anomalous m3 with 15 plates which correlates with the Bavel Interstadial of an advanced M. meridionalis was dredged (the first interstadial of the Bavelian) accor- from the floodplain of the river Meuse near ding to Zagwijn & De Jong (1984). This age Alphen aan de Maas (Province of Gelder- would be in accordance with the advanced land). The stratigraphical position is un- evolutionary stage of the M. meridionalis known but could be late Early to early M i d d l e molars from Dorst-Surae.

126 MOL et al.: Netherlands proboscideans

Figure 4 Metacarpus III dex. of A . a r v e r n e n s i s (A C o l l . NNM no. St-119327) and M . m e ri d i o n a l i s (B C o l l . NNM no. S t - 1 4 5 5 8 5 ) , both dredged up from the Oosterschelde Estuary and both shown in anterior view. Note the steeper inclination of the facets fo r a r ticulation with the magnum and uncinatum and the wider protrusion of the distal and proximal epiphyses in A n a n c u s.

From the same pit at Dorst, Van Kolfschoten cm), the lamellar frequency of 5.1, and the (1990) described one m3 (Coll. NNM, St- enamel thickness (1.9-3.1 mm) all fall within 85531) and attributed this to Elephas anti - the range of a m2 of M. meridionalis a c c o r- q u u s. Our examination of St-85531 revealed ding to Maglio (1973). that Van Kolfschoten misidentified the molar: it is not an m3 but a very worn-down remnant Lister (1993) places the M. meridionalis/M. of a left m1 or m2. The maximum width (7.2 t ro g o n t h e r i i boundary at 0.7-0.6 My. T h e

127 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

Dorst-Surae M. meridionalis are probably the Some superb E. antiquus remains are found youngest dated fossils of this species in T h e in the collections of the Koninklijk Zeeuwsch Netherlands, though some of the North Sea Genootschap der Wetenschappen, stored in material may be younger or equivalent in age the Zeeuws Museum at Middelburg. One with the Dorst-Surae mammoths. l a rge hemi-mandible with the m3, and some extremely large, heavily built postcranial Elephas antiquus remains, which were trawled by fisherman The straight-tusked elephant, Elephas anti - from the mouth of the Westerschelde, are q u u s, is well known from temperate stored in the collections. From the same European interglacials, especially from the Schelde Estuary a very large humerus, also late Middle and Late Pleistocene. Several heavily built, attributed to E. antiquus, is finds are recorded from The Netherlands and stored in the NNM at Leiden. Again from the the North Sea. Their geological age is not Westerschelde, the Late Pleistocene always clear. Most of the finds are from Hippopotamus incognitus is known. A L a t e below the water level of the sea or from dred- Pleistocene age is inferred, based on the ging operations alongside the great rivers, occurrence of the latter species and on the e.g. the rivers Rhine and IJssel. Roding state of fossilization of the We s t e r s c h e l d e (1953) described some molar fragments of E . material. The material is characterised by a a n t i q u u s from the Needse Berg (Neede, low density and weak mineralisation. Province of Gelderland). The scarce remains, which have been found in situ, originate from The Eemian fauna of Haerst (Province of Middle Pleistocene sediments, which are Overijssel) includes, amongst others, E. anti - placed in the Holstein Interglacial. A c c o r d i n g q u u s and H. incognitus. Again from this site, to Van Kolfschoten (1988) it is not clear if sampled by a dredging operation, only pro- the elephant remains from the Neede site boscidean molars are known of the straight- belong to E. antiquus or M. meridionalis. tusked elephant. It is the northernmost occur- One of the fragments is rather brachyodont. rence of E. antiquus in The Netherlands. Van Kolfschoten (1981) mentioned some molar fragments of E. antiquus from the Hooijer (1984) described a fragment of Middle Pleistocene ice-pushed ridge near E. antiquus (then called E. namadicus) that Rhenen (Prov. of Gelderland). From the i n was trawled from the bottom of the North s i t u site Belvédère near (Province Sea. Since then more material has been col- of Limburg), Van Kolfschoten (1988) mentio- lected. This material is stored in private col- ned the straight-tusked elephant of lections. So far one mandible with an m3 and Maastricht-Belvédère-2 and -4, which are four isolated molars of E. antiquus have been placed in the early Saalian (late Middle recognised from the Southern Bight of the P l e i s t o c e n e ) . North Sea. The state of preservation is the A molar of E. antiquus is known from dred- same as usual in the Late Pleistocene terres- ging operations in the floodplain of the river trial mammal remains from the North Sea IJssel near Giesbeek (Province of Gelder- and the Westerschelde, that is, low density land). It originates probably from the same and weak mineralisation. The geological age sediments in which the Late Pleistocene is unknown. However, we suggest a late Hippopotamus incognitus was recovered Middle or Late Pleistocene age. If the (Mol 1993). From the Maasvlakte, an artifi- remains of E. antiquus from the North Sea cial peninsula on the coast of the Province of originate from Eemian deposits, they might Zuid-Holland, E. antiquus is mentioned by indicate a land connection of the British Isles Mol (1994). Mol (1994) suggests a Late with the continent. E. antiquus is well known Pleistocene (Eemian) age for E. antiquus. from various sites in the British Isles and the

128 MOL et al.: Netherlands proboscideans

Figure 5 The same specimens as in Figure 4, here shown in lateral view. Note the relative ly more prox i m a l ly extending distal j o i n t s , the steeper proximal articulation facet and the more anteroposteri o r ly protruding epiphyses in A n a n c u s (A) as compared to M a m m u t h u s (B) .

continent with an Eemian age (Sutcliffe 1985). of Dorst-Surae. There is no evidence that As already noted above, the molar from E. antiquus was already present in T h e Dorst-Surae, described by Van Kolfschoten Netherlands during the Early Pleistocene. (1990: fig. 6) as E. antiquus, with an Early Pleistocene age (Bavelian), should be attribu- M a m muthus primigenius ted to M. meridionalis. Therefore, E. anti - Proboscidean remains most frequently reco- q u u s should be removed from the fauna list vered from the North Sea bottom are those of

129 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

the woolly mammoth (Mol 1989, 1991). The Netherlands is that of a male skull of an Thousands of fossils have been collected so adult individual. Age assessment according to f a r. Van Essen & Mol (1996) give measure- the dental criteria set up by Laws (1966) ments of some dental elements of M. primi - gives an estimated age of 43 ± 2 A E Y f o r genius for comparison with the M. meridio - this individual. 1 4C dating carried out at n a l i s teeth. M. primigenius is amongst the Utrecht University gave an absolute age of most abundant species of the We i c h s e l i a n 31,000 ± 400 yBP (UtC-4551). This skull Mammoth Fauna from the North Sea. T h i s was found after a flooding of the River Linge fauna includes: Homo sapiens, Castor fiber, near Heukelum (Province of Zuid-Holland) Canis lupus, Vulpes vulpes, Ursus spelaeus, in 1820. The skull was described by Va n Ursus arc t o s , C rocuta crocuta spelaea, Marum (1824) and by Cuvier (1834). T h e Panthera leo spelaea, Mammuthus primigeni - history of this early find, which is now in the u s, Equus caballus, Equus hydru n t i n u s , Teylers Museum at Haarlem, was described Coelodonta antiquitatis, Sus scro f a, by Mol et al. (1995, 1996). In the same paper M e g a l o c e ros giganteus, Alces alces, C e rv u s most of the skulls of woolly mammoth stored e l a p h u s , C a p reolus capre o l u s, R a n g i f e r in Dutch museums are listed. Two are well t a r a n d u s, Ovibos moschatus, Bison priscus preserved and have been 1 4 C dated. and Bos primigenius. The first one is a superb skull with tusks and Mammal remains attributable to these taxa mandible with complete dentition. It was dred- originate from the Brown Bank or Brown ged from a depth of 12 m below the water Ridge, and are characterized by a low density level in the floodplain of the River IJssel near and weak mineralisation. The state of preser- O l b u rgen (Province of Gelderland). The skull vation is in most cases excellent. Most of the belonged to a 40 A E Y old and very small material is winnowed from the seabed by female individual. Its estimated shoulder marine currents, and depending on the time height is less than 250 cm. A 1 4C age of they were lying on the sea bottom the fossils 22,160 ± 260 yBP (UtC-4550) was obtained are overgrown to various degrees with bryo- for this specimen. zoans and barnacles. Unfortunately, to date, there are no 1 4C datings available of the Late The second specimen is a nice, slightly dama- Pleistocene terrestrial mammal material from ged cranium, the mandible missing, of an old the North Sea. Some of the above listed spe- bull, which was brought to daylight by divers cies therefore could have an Early Holocene in the artificial lake ‘De Groene Heuvels’ n e a r age (Van Kolfschoten & Laban 1995). T h e B o rgharen (Province of Gelderland) in 1994. extensive collection of M. primigenius m a t e- The 1 4 C dating results for this specimen are rial (the largest collection is to be found in 38,900 ± 900 yBP (UtC-3621, Thijs van the NNM) shows that all ontogenetic stages Kolfschoten, personal communication 1996). are represented and also that the size variabi- An accumulation of many hundreds of lity is considerable. Some of these remains Weichselian woolly mammoth bones was are very small and may represent the latest recovered in the 1960's in the floodplain of the populations before this species became River Meuse near Gewande, north of 's- extinct in Western Europe. Hertogenbosch (Province of Noord-Brabant). The Gewande collection, which belongs to M. primigenius remains have also been reco- one of the best in The Netherlands, was stored vered from many sites onshore, though larg e in the Institute of Earth Sciences of Utrecht bone accumulations like those occurring in U n i v e r s i t y, and was recently moved to the Siberia are not known from The Netherlands. NNM (Naturalis). Unfortunately, the collec- One of the earliest finds of M. primigenius i n tion was never studied in detail. A partial ske-

130 MOL et al.: Netherlands proboscideans

Figure 6 A M e t a c a r pus V sin. of Mammuthus meri d i o n a l i s ( C o l l . NNM no. S t - 1 1 9 0 4 4 ) , l a t e ral view. B M e t a c a r pus II sin. o f Mammuthus meri d i o n a l i s ( C o l l . NNM no. S t - 1 4 5 9 0 1 ) , distal view. A n t e rior side is up. Note the pointed profile of the distal a r ticulation surface (A) and the presence of distinct fovea on this surface (B) .

131 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

leton is known from Gewande. About 27 silization, as well as a lower m1 and a frag- remains, mainly large bones and some footbo- mentary m2, is assigned to a female individu- nes, were attributed by Mol (1984) to one and al. The molars indicate an age of about 15 the same individual, a relatively young fema- A E Y. A fragmentary humerus was attributed le. A well preserved skull of a large old male to a juvenile individual with a shoulder height from Gewande was published and figured by of approximately 90 cm. To date, the Orvelte Mol (1992) when he compared it with another fossils are the only in situ mammoth remains interesting, pathological skull of a woolly from The Netherlands that have been 1 4 C mammoth. This latter skull, including the dated. The dating results are 46,800 +1500/- mandible, was dredged from a gravelpit at 1250 yBP (GrN-18780, Van der Sanden 1993). Va l b u rg (Province of Gelderland). The molars in the upper and lower jaw (M3 sin. and dex. The oldest M. primigenius remains in T h e and the m3 dex.) show that we are dealing Netherlands are known from a sand pit called with a skull of an old individual with an esti- ‘De Fransche Kamp’ at Wa g e n i n g e n mated age of 53 ± 2 A E Y. The dentition is (Province of Gelderland). According to Va n anomalous: the m3 in the left lower jaw is Kolfschoten (1988) the mammoths from this missing. As its alveole is worn and the crown site should be placed in the Early Saalian and of the M3 sin. is considerably higher than that might have an age of approximately 250,000 of the M3 dex. (the difference is 110 mm) it y e a r. can be deduced that the left m3 was lost i n v i v o. At Maastricht-Belvédère-2, M. primigenius i s found together with E q u u s sp., C o e l o d o n t a In situ sites of M. primigenius are very rare in a n t i q u i t a t i s and C e rvus elaphus ( Va n The Netherlands. Van der Meulen (1991) Kolfschoten 1988). A nearly complete curved described remains that were found in a con- and spirally twisted tusk with a length of struction pit near Grou (Province of Friesland) three metres and some molars from the same in the northeastern part of The Netherlands. specimen were collected in situ. According to Remains of Pleistocene mammals are very Van Kolfschoten (1988), Maastricht- rare in the northern provinces. According to Belvédère-2 is of an Early Saalian age, pro- Boekschoten (1965) they are almost complete- bably the same as that of Wa g e n i n g e n - ly absent in the Saalian and Weichselian depo- ‘Fransche Kamp’. The Maastricht-Belvédère- sits of the northern Netherlands. The in situ 2 mammoths lived in a tundra-steppe envi- finds of Grou were situated in Early ronment with cool climatic conditions (Va n Weichselian sediments, deposited in shallow Kolfschoten 1988). Maastricht-Belvédère-5, fresh water. of an Early Weichselian age, also has M. pri - m i g e n i u s together with woolly rhino in its Remains of woolly mammoths were excavated fauna. According to Van Kolfschoten (1988) at Orvelte (Province of Drenthe) during a res- the Maastricht-Belvédère-5 fauna composi- cue excavation covering 80 square metres. tion points to a tundra-steppe environment The remains represent at least three, and per- and a cold and rather dry climate during the haps four individuals. Most of the excavated period in which the fauna lived. material, amongst others the mandible with both m3’s and many cranium fragments, was C O M PARISON OF MANUS AND PES attributed to one single individual: a bull BONES OF A N A N C U S AND approximately 46 A E Y old and with an esti- M A M M U T H U S mated shoulder height of 280 cm (Mol & Va n Postcranial parts attributed to A. arv e r n e n s i s Kolfschoten 1993, Van Kolfschoten & Mol are considerably smaller than those of most 1993). The tip of a tusk, broken off before fos- other Pleistocene and recent proboscideans

132 MOL et al.: Netherlands proboscideans

Ta ble 1 Size measurements (in mm) of metacarpalia of A . a r v e r n e n s i s and M . m e ri d i o n a l i s from the Oosterschelde Estuary and the N o r th Sea, present in the NNM collection and the Collection Mol.The measurements are defined in Appendix 1(1).

(excluding the pygmy from the total body length (excluding tusks) of 4.2 m. Mediterranean islands). A complete A. arv e r - The A. arv e r n e n s i s molar remains from the n e n s i s skeleton was found in 1826 by Filipo Oosterschelde seem particularly small when Nesti near Montecarlo (Val d'Arno, Tu s c a n y, compared with specimens cited by To b i e n ) and is now exhibited in the Geological (1973). Schreuder (1944) also reached this and Paleontological Museum in Florence. conclusion after comparison with data from Osborn (1936: fig. 595) presented a 1:100 the older literature. The elements of manus scaled reconstruction of A. arv e r n e n s i s. He and pes are particularly suited to demonstrate gave a shoulder height of 2550 mm and a the relatively small size, as they are often

133 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

Ta ble 2 Size measurements (in mm) of carpalia of A . a r v e r n e n s i s and M . m e ri d i o n a l i s from the Oosterschelde Estuary and the N o r th Sea, present in the NNM collection and the Collection Mol.The measurements are defined in Appendix 1 (2-7).

found intact. The collection of the NNM con- tions, except for the trapezium. In addition, tains the following elements from the there are one or more M. meridionalis s p e c i- Oosterschelde that could be attributed to mens of the following elements in the same A. arv e r n e n s i s: a juvenile right metacarpus II collections: metacarpus IV, lunatum, uncina- lacking the distal epiphysis (no. St-118412); a tum, pisiforme, calcaneus and naviculare. right metacarpus III (no. St-119327); a left metacarpus V (no. St-118902); a right trape- M. meridionalis was one of the largest pro- zium (no. St-401477); a left trapezoideum boscideans from the Eurasian Pleistocene (no. St-401409), and a right astragalus (no. with a shoulder height of about 4 m. T h e St-140740). Furthermore, there is a left mag- Nogaisk skeleton in the St. Petersburg num of A. arv e r n e n s i s originating from the Zoological Museum stood 420 cm at the Thornton Bank in the collection Mol (no. s h o u l d e r, that of Georgyevsk in the Stavropol 2 0 0 6 ) . Museum 396 cm. The maximum lengths of the femora are 146 and 143.5 cm respectively M. meridionalis hand and foot bones are (Garutt 1964). Diverse large-sized skeletal more frequently dredged from the Ooster- parts from East Anglia (Britain) were descri- schelde and the North Sea. M. meridionalis bed by Adams (1877-1881). A c o m p l e t e equivalents of the available A. arv e r n e n s i s femur from Mundesley has a maximum elements are in the NNM and Mol collec- length of 150 cm (Adams 1881). An excep-

134 MOL et al.: Netherlands proboscideans

Ta ble 3 Size measurements (in mm) of tarsalia of A . a r v e rn e n s i s and M . m e ri d i o n a l i s from the Oosterschelde Estuary and the N o r th Sea, present in the NNM collection and the Collection Mol.The measurements are defined in Appendix 1(8-10).

tionally complete skeleton from Scoppito, Sangerhausen (Germany) stands about 250 l'Aquila (Italy) was described by Maccagno cm at the shoulder (Krutsch 1953; Garutt & (1962) and stood 400 cm at the shoulder. T h e Nikolskaja 1988). A male skeleton in the likewise very complete skeleton from Borro Azov Museum reaches 450 cm at the shoul- al Quercio, Arezzo (Valdarno, Italy) measures der according to Garutt & Nikolskaja (1988). 380 cm at the shoulder. It must be noted that This height may be slightly overestimated it is doubtful whether skeletal parts of because the vertebrae have been mounted M. tro g o n t h e r i i can be distinguished from between the tips of the shoulderblades and those of M. meridionalis. The basis for com- therefore the dorsal spines protrude too much parison is relatively small. The nearly com- above the shoulderblades. A humerus from plete M. tro g o n t h e r i i skeleton from Mosbach Sande near Wiesbaden (Germany) Edersleben in the Spengler Museum at is 144 cm long, 15 cm longer than the hume-

135 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

rus in the Azov skeleton. Since the presence our case the large size difference between the of M. trogontherii in the Southern Bight of A n a n c u s and M a m m u t h u s elements facilitates the North Sea was demonstrated by several their distinction. As follows from the meas- teeth, there is a possibility that some postcra- urements given in Tables 1-3, the A. arv e r - nial parts from the North Sea referred to here n e n s i s bones in the studied sample are on as M. meridionalis, were misinterpreted. A l s o average between 60 and 70 percent smaller sporadic E. antiquus molars have been dredg- than their equivalents in M. meridionalis. ed from the Southern Bight of the North Sea, Apart from their small size, the metacarpals, but their weak degree of fossilization diff e r s carpals and tarsals of A. arv e r n e n s i s a r e from the heavily mineralized bones here attri- generally speaking characterised by their well buted to M. meridionalis. delineated and more concave/convex or more Tables 1-3 give the size measurements of the inclined proximal and distal articulation various A. arv e r n e n s i s and M. meridionalis facets, when compared to M. meridionalis metacarpals, carpals and tarsals studied. (Fig. 4). Carpals and tarsals of the latter spe- Unless otherwise stated, the specimens have cies have overall flatter articulation facets on been dredged from the Oosterschelde. T h e the proximal and distal surfaces, better suited definitions of the measurements taken are for the transfer of force but allowing only indicated in Appendix 1 (1-10). Not included limited movement. Furthermore, the metacar- in the tables are specimens of M a m m u t h u s pals of A n a n c u s have comparatively widely p r i m i g e n i u s from the North Sea. These light- protruding distal and proximal terminations, ly mineralized fossils are usually intermediate as follows from the relatively low MC1/MC3 in size. Material of the woolly mammoth ratios, but similar MC1/MC6 ratios as com- from the North Sea is much more numerous pared to M. meridionalis ( Tables 1b and 1d). and will be dealt with in another paper. Of course, the material available to us, espe- Intraspecific variation in the shape of probos- cially that of A. arv e r n e n s i s, is too limited to cidean (meta)carpals and (meta)tarsals is rat- assess intraspecific variation and overlapping her large. Especially the morphological out- morphologies, so that at the moment these line of the articulation facets is quite variable. remarks cannot be more than general state- Considering the metapodials, the ratio ments. However, there are a few characteris- between length and transverse or anteroposte- tic differences that should be noted. First of rior diameter seems subject to considerable all, the distal articulation surface of the A . variation. Compare, for example, the ratios a rv e r n e n s i s metacarpals continue more proxi- between various measurements of the three mally on the anterior and posterior sides. In metacarpals-II attributed to M. meridionalis sagittal cross section these articulation sur- in Table 1a: especially NNM St-145341 is faces describe an almost perfect halfcircle, quite slenderly built as expressed by the rela- whereas in M. meridionalis metacarpals the tively high MC1/MC3 and MC1/MC6 ratios. cross-sectional outline corresponds with a A similar wide range in the length/width segment less than twofifth of a circle (Fig. 5). ratios is observed in the three metacarpals-V This difference is clearly expressed by the attributed to M. meridionalis ( Table 1d). T h e small to very small ratio in A n a n c u s b e t w e e n relatively large morphological variation is total metacarpal length and the vertical height probably due to the fact that ontogenic deve- of the articulation facet (ratio MC1/MC9, lopment is much constrained by the space left Tables 1b, 1d). Also, the articulation facet is available by the surrounding bones. The larg e not perfectly round but rather pointed in most intraspecific morphological variation often Mammuthus metapodials (Fig. 6). Obviously, hampers the determination of isolated ele- A n a n c u s metacarpals stood more closely to ments to species and even to level. In the ancestral condition where the hinging

136 MOL et al.: Netherlands proboscideans

movement between metapodials and digits from the Southern Bight of the North Sea and could take place freely, whereas in gravipor- the Westerschelde Estuary. Proboscidean tal limbs of M. meridionalis hinging move- material dredged or excavated from sand and ment had been largely reduced. Another dis- gravel pits located on the Dutch alluvial tinction can be made at the proximal articula- plains are less frequently encountered, though tion surfaces of the metacarpals II and III. the stratigraphic context is usually better The anteroposterior crest dividing the proxi- documented. A few mainland localities have mal articulation surfaces into two facets (for yielded M. primigenius remains from single the trapezoideum and magnum in the individuals. Commonly dredged elements of Metacarpus II and for the magnum and unci- fossil proboscideans which are complete are natum in the Metacarpus III) is clearly shar- the (meta)carpals and tarsals. These elements per in A. arv e r n e n s i s (Fig. 4). Especially the can be well distinguished in the case of narrow lateral facets for articulation with the A. arv e r n e n s i s and M. meridionalis, due the magnum and uncinatum, respectively, are size difference but also on the basis of mor- more inclined in the latter species. The same phological grounds. holds true for the corresponding medial facet on the distal articulation surface of the mag- AC K N OW L E D G E M E N T S num, which articulates with the metacarpus II We thank Dr. K. van der Borg (Utrecht (no uncinatum of A. arv e r n e n s i s is present in University) for providing the 1 4C dating the collections). The foot construction of results of the Heukelum and Olburger woolly Anancus arv e r n e n s i s seems to have allowed mammoth skulls. We also want to thank more internal movement, and was probably numerous private collectors and members of better adapted to walk over soft soils than the museum staffs in The Netherlands for making graviportal limbs of M. meridionalis. fossil proboscideans in their collections avai- lable for this review. We are especially indeb- C O N C L U S I O N S ted to Mr. Cor Strang of Naturalis (Leiden) L a rge numbers of proboscidean and other ter- who helped us in many ways during the pre- restrial mammals have been brought to light paration of this paper. The photographs were as a by-product of trawling in the Southern taken by Mieke van Engelen. Reinier van Bight of the North Sea and the Oosterschelde Zelst prepared the plates. E s t u a r y. The oldest fauna association from The Netherlands, with Anancus arv e r n e n s i s R E F E R E N C E S and Mammuthus meridionalis originates from Adams, A. Leith, 1877 - Monograph on the British Fossil the Oosterschelde Estuary and has an estima- Elephants, Volume I - The Palaeontographical ted age of around 1.9 My, slightly older than S o c i e t y, London the 1.7 My old Tegelen fauna. A few molars Adams, A. Leith, 1879 - Monograph on the British Fossil of Mammut borsoni from Liessel may be Elephants, Volume II - The Palaeontographical older in age than the Oosterschelde assembla- S o c i e t y, London ge. A heavily fossilized Early to Middle Adams, A. Leith, 1881 - Monograph on the British Fossil Pleistocene fauna association originates from Elephants, Volume III - The Palaeontographical the North Sea, mainly from the Deep Wa t e r S o c i e t y, London Channel. This material includes the probosci- Azzaroli, A. & Mazza, P., 1992 - The cervid genus deans M. meridionalis and rare M. tro g o n t h e - E u c l a d o c e ro s in the early Pleistocene of Tuscany - r i i fossils. A Late Pleistocene fauna associa- Palaeontographia Italica 79: 43-100 tion including abundant remains of M. primi - Azzaroli, A., De Giuli, C., Ficcarelli, G. & Torre, D., g e n i u s originates from the Brown Bank and 1988 - Late Pliocene to early Mid-Pleistocene has a light degree of fossilization. Also a few mammals in Eurasia: faunal succession and dispersal lightly fossilized E. antiquus fossils originate events - Palaeogeography, Palaeoclimatology,

137 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

Palaeoecology 66: 77-100 m e r i d i o n a l i s und von Trogontherium cuvieri aus den Boekschoten, G.J., 1965 - Over de verspreiding van I n t e rglazialen Tegelen-Schichten - Quartär- mammoetfossielen in noordelijk Nederland - paläontologie 6: 53-65 Geologie en Mijnbouw 44: 295-297 H o o i j e r, D.A., 1950 - Het belang van de vertebraten voor Boeuf, O., 1983 - Le Site Villafranchien de Chilhac de stratigraphie van het Nederlandse Kwartair ouder (Haute-Loire) France. Etude Paléontologique et dan de ijsbedekking - Geologie en Mijnbouw, new Biochronologique - Unpublished Ph.D. dissertation, series 12: 12-13 Université Paris V I H o o i j e r, D.A., 1953 - On dredged specimens of A n a n c u s , Boeuf, O., 1993 - Il était une fois. Il-y-a près de A rc h i d i s k o d o n and E q u u s from the Schelde estuary, 2.000.000 d'années a Chilhac, Haute-Loire, France - Netherlands - Leidse Geologische Mededelingen 17: Amis de Chilhac (Association): 1-49 1 8 5 - 2 0 1 Bout, P., 1970 - Absolute ages of some volcanic forma- H o o i j e r, D.A., 1957 - Mammals and correlation of the tions in the A u v e rgne and Velay areas and chronology ‘Black Bones’ of the Schelde estuary with those of of the European Pleistocene - Palaeogeography, the Red Grag - Geologie en Mijnbouw, new series P a l a e o c l i m a t o l o g y, Palaeoecology 8: 95-106 19: 255-256 Bout, P., 1979 - Sédimentation villafranchienne, volca- H o o i j e r, D.A., 1962 - The sabre-toothed cat nisme et tectonique du versant Est de l'extrémité H o m o t h e r i u m found in The Netherlands - Lutra 4 (1): septentrionale du Devès - Publication de l'Institut de 24-26 Géographie de la Faculté des Lettres de Clermont- H o o i j e r, D.A., 1981 - The first rhinocerotid of Ferrand 57: 25-32 the Pretiglian black bones from the Netherlands - Bout, P. & Azzaroli, A., 1952 - Stratigraphie et faune de Netherlands Journal of Zoology 31 (2): 472-475 Creux de Peyrolles près Perrier (Puy-de-Dôme) - H o o i j e r, D.A., 1984 - Mammuthus meridionalis (Nesti) Annales de Paléontologie 38: 35-56 and M. armeniacus (Falconer) from the North Sea - Cameron, T.D.J., Laban, C. & Schüttenhelm, R.T.E., Proceedings of the Koninklijke Nederlandse 1984 - Flemish Bight, Sheet 52 -02 E. Quaternary Akademie voor Wetenschappen, Series B, 87: Geology 1:250,000 - British Geological Survey and 3 3 5 - 3 5 9 Rijks Geologische Dienst, Southampton H o o i j e r, D.A., 1991 - Fossielen vissen voor de C u v i e r, G., 1834 - Recherches sur les Ossemens Fossiles, wetenschap; veertig jaar Kor en Bot (1951-1991) - Tome II - Quatrième Editions, Paris:1-500 K r o n i e k van het Land van de Zeemeermin De Vos, J., Mol, D. & Reumer, J.W. F., 1995 - The early (Schouwen-Duiveland): 1-16 Pleistocene cervids of the Oosterschelde (The Kortenbout van der Sluijs, 1985 - Botten uit de Netherlands) and a revision of the cervid genus Oosterschelde - Cranium 2 (1): 9-10 E u c l a d o c e ro s F a l c o n e r, 1868 - Deinsea 2: 95-123 Krutsch, W., 1953 - Ein eiszeitliches Elefantenskelett im Drees, M., 1986 - Kritische kanttekeningen bij de naam S p e n g l e r-Museum Sangerhausen - Natur und ‘Zwarte Botten Fauna’ - Cranium 3 (1): 103-120 Museum 4: 11 4 - 11 6 Dumon Tak, A.M., 1973 - Werkgroep Palaeontologie Laws, R.M., 1966 - Age criteria for the , ‘ B o t t e n v i s t o c h t ’ op de Westerschelde - Zeeuws Loxodonta a. africana - East African Wildlife Journal Tijdschrift 23(4): 21-22 4: 1-37 Garutt, V.E., 1964 - Das Mammut M. primigenius L i s t e r, A.M., 1993 - ‘Gradualistic’ Evolution: its (BLUMENBACH) - A. Ziemsen Verlag, Wi t t e n b e rg Interpretation in Quaternary Large Mammal Species - L u t h e r s t a d t Quaternary International 19: 77-85 Garutt, V.E. & Nikolskaja, V.N., 1988 - Über das Skelett Maccagno, A.M., 1962 - L'Elephas meridionalis Nesti di vom Steppenelefanten aus Edersleben - Spengler Contrada ‘Madonna della Strada’ Scoppito (L'Aquila) Museum (Sangerhausen, Germany) 9: 3-13 - Atti dell’ Accademia delle Scienze fisische e mate- Germonpré, M., 1983 - Les mammifères de la Formation matische di Napoli III, 4, 1: 1-132 de la Campine - Bulletin Belgische Vereniging voor Maglio, V.J., 1973 - Origin and evolution of the Geologie 92(2): 111-123 - Transactions of the American G u e n t h e r, E.W., 1986 - Funde von A rc h i d i s k o d o n Philosophical Society 63: 1-149

138 MOL et al.: Netherlands proboscideans

Mol, D., 1984 - Postcraniale skeletdelen van een indivi- Firenze nel 1841: 159 du van de mammoet uit Nederland - Cranium 1 (2): Osborn, H.F., 1936 - Proboscidea. A monograph of the 4 7 - 4 9 d i s c o v e r y, evolution, migration and of the Mol, D., 1989 - Fossil proboscidea from the North Sea mastodonts and elephants of the world. Volume 1: 1- bottom between England and The Netherlands - 802 - Published on the J. Pierpont Morgan Fund by I . T.C. congress, Rome 22-29 August 1989, Abstracts the Trustees of the American Museum of Natural of papers and posters, Vol. I: 154-155 H i s t o r y, The American Museum Press, New Yo r k Mol, D., 1991 - Het IJstijdlandschap van de zuidelijke Peters, A., Lammers, Th. & Mol, D., 1991 - Noordzee - Grondboor en Hamer 45 (1): 9-13 Mastodonten-kiezen uit Liessel (Noord-Brabant) - Mol, D., 1992 - Een bijzondere mammoetschedel uit Cranium 8: 89-96 Va l b u rg - Grondboor en Hamer 46 (1): 18-23 Roding, G.M., 1953 - Elephas antiquus uit Neede - Mol, D., 1993 - Nijlpaarden dobberden in de IJssel - Publicatie 14 van de Nederlandse Geologische Grondboor en Hamer 47 (3): 73-79 Ve r e n i g i n g Mol, D., 1994 - Nog meer nijlpaarden uit Nederlandse Rutten, L.M.R., 1909 - Die diluvialen Säugetiere der bodem - Grondboor en Hamer 48 (1): 7-8 Niederlande - Ph.D. Dissertation, University of Mol, D. & Van Essen, J.A., 1990 - Mammut borsoni Utrecht, J.van Boekhoven Press, Utrecht from The Netherlands - Lutra 33: 183-186 S c h r e u d e r, A ., 1944 – Upper Pleistocene Proboscidea out Mol, D. & H. Van Essen, 1992 - De Mammoet, sporen of the Scheldt and the Lower Rhine - Leidse uit de IJstijd - Uitgeverij BZZTôH, ‘s-Gravenhage: Geologische Mededelingen 14: 40-58 1 - 1 3 9 S c h r e u d e r, A ., 1945 - De mastodont en de olifant onder Mol, D. & Van Kolfschoten, Th., 1993 - De mammoeten de ‘zwarte fossielen’ uit de Zeeuwsche wateren - van Orvelte en andere dieren uit de ijstijd - in: Van Verhandelingen van het Geologisch en der Sanden, W.A.B., Cappers, R.T.J., Beuker, J.R. & Mijnbouwkundig Genootschap 14: 437-448 Mol, D. (eds.) - Mens en Mammoet. De mammoeten Spaan, A., 1992 - A revision of the deer from Tegelen van Orvelte en de vroegste bewoning van Noord- (province of Limburg, The Netherlands) - Scripta Nederland - Archeologische Monografieën van het Geologica 98: 1-85 Drents Museum 5: 14-26 S u t c l i ffe, A.J., 1985 - On the track of ice age mammals Mol, D. & De Vos, J., 1995a - Korren op de - British Museum (Natural History), London Oosterschelde; een zoogdierpaleontoloog als visser Tobien, H., 1973 - The Structure of the Mastodont Molar en wat de fossielen van de Oosterschelde ons (Proboscidea, Mammalia). Part 1: The Bunodont vertellen - Grondboor en Hamer 49 (3/4): 57-61 Pattern - Mainzer geowissenschaftlichen Mol, D. & De Vos, J., 1995b - De hyena uit de Mitteilungen 2: 11 5 - 1 4 7 Oosterschelde - Grondboor en Hamer 49 (6): 139-149 Van der Feen, P.J., 1968 - A fossil skull fragment of a Mol, D. & De Vos, J., 1996 - Early Pleistocene mamma- walrus from the mouth of the river Scheldt lian remains from the Eastern Scheldt (Zeeland (Netherlands) - Bijdragen tot de dierkunde 38: 23-30 Province, The Netherlands) – in: Volume of A b s t r a c t s Van der Meulen, S., 1991 - Mammoetresten of the Inqua-SEQS, ‘The dawn of the Quaternary’, (Mammuthus primigenius) uit een in situ positie in 16-21 June, 1996, Kerkrade, The Netherlands: 71 het noordoosten van Nederland (Grou, Friesland) - Mol, D., Ter Mors, G., Van Veen, J., & De Vos, J., 1995 - Cranium 8 (2): 63-64 De geschiedenis van de mammoetschedel van Van der Sanden, W.A.B., 1993 - Het verhaal van een Heukelum - Teylers Magazijn 49: 9-14 t o e v a l s t r e ffe r - in: van der Sanden, W.E.B., Cappers, Mol, D., Ter Mors, G., Van Veen, J., & De Vos, J., 1996 - R . T.J., Beuker, J.R. & Mol, D. (eds.) - Mens en De geschiedenis van de mammoetschedel van Mammoet. De mammoeten van Orvelte en de vroeg- Heukelum en iets over mammoeten en hun schedels - ste bewoning van Noord-Nederland - Archeologische Grondboor en Hamer 50: 17-23 Monografieën van het Drents Museum 5: 7-13 Nesti, F., 1841 - Report of the meeting of Sept. 25t h , Van der Vlerk, I.M., 1951 - Zeeland in het IJstijdvak - summarized by P. Savi and A. Sismonda - Atti della Koninklijke Nederlandse Akademie voor Terza Riunione degli scienziati Italiana tenuta in Wetenschappen, Akademiedagen 4: 11 0 - 1 2 4

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Van Essen, H. & Mol, D., 1996 - Plio-Pleistocene pro- Van Kolfschoten, Th. & Laban, C., 1995 - Pleistocene boscideans from the southern bight of the North Sea terrestrial mammal faunas from the North Sea - and the Eastern Scheldt, The Netherlands - in: Mededelingen Rijks Geologische Dienst 52: 135-151 Shoshani, J. & Ta s s y, P. (eds.) - The Proboscidea: Van Marum, M., 1824 - Verhandeling over de Olijfanten, Trends in Evolution and Palaeoecology – pp. 214- die in vroeger eeuwen de noordelijke gematigde en 224. Oxford University Press koude streken der aarde bewoonden, waarin eene Van Kolfschoten, M., 1981 - On the Holsteinian? and beschrijving en afbeelding van de olijfantskop, welke Saalian mammal fauna from the ice-pushed ridge in 1820 uit een kuil nabij Heukelum, bij doorbraak near Rhenen (The Netherlands) - Mededelingen van ontstaan, is uitgespoeld – Natuurkundige verhandelin- de Rijks Geologische Dienst 35(6): 223-251 gen Hollandsche Maatschappij der Wetenschappen Van Kolfschoten, M., 1988 - The evolution of the 13: 253-304 mammal fauna in The Netherlands and the Middle Von Koenigswald, G.H.R., 1950 - Voorlopige mede- Rhine Area (Western Germany) during the late deling omtrent het voorkomen van borsoni Middle Pleistocene - Unpublished Ph.D. Thesis, in Nederland - Geologie en Mijnbouw, Nieuwe Serie Utrecht University 12: 14-15 Van Kolfschoten, Th., 1990 - The Early Biharian Zagwijn, W.H., 1960 - Aspects of the Pliocene and Early Mammal Faunas from Bavel and Dorst-Surae - Pleistocene Vegetation in The Netherlands - Ph.D. Quartärpaläontologie 8: 265-272 Dissertation, Leiden State University, Leiden Van Kolfschoten, Th. & Van der Meulen, A.J., 1986 - Zagwijn, W.H., 1963 - Pollen-analytic investigations in Villanyian and Biharian mammal faunas from the the Tiglian of The Netherlands - Mededelingen van de Netherlands - Memorie Società Geologica Italiana 31: Geologische Stichting, Nieuwe Serie 16: 49-71 1 9 1 - 2 0 0 Zagwijn, W.H. & De Jong, J., 1984 - Die Interglaziale Van Kolfschoten, Th. & Mol, D., 1993 - De mammoeten von Bavel und Leerdam und ihre stratigraphische en de neushoorn van Orvelte (Drenthe) - Cranium 10 Stellung im niederländischen Frühpleistozän - Mede- (2): 101-111 delingen van de Rijks Geologische Dienst 37: 155-1 6 9

140 MOL et al.: Netherlands proboscideans

APPENDIX 1 D e finition of the size measurements taken on the metacarp a l i a , c a rpalia and tarsalia of Ta bles I-3. DT = tra n s ve rse diameter, DAP = anteroposterior diameter.

1 M e t a c a r p a l s . A = anterior view of metacarpus III dex.; B = lateral view of metacarpus III dex.; C = posterior view of m e t a c a rpus V sin. MC1 = Maximum length; MC2 = Length between the articulation surfaces (= MC1 except for the m e t a c a rpus V ) ; MC3 = DT prox i m a l ; MC4 = DAP prox i m a l ; MC5 = Minimum DT diaphy s i s ; MC6 = Minimum DAP diaphy- s i s ; MC7 = Maximum distal DT; MC8 = Maximum distal DA P ; MC9 = Height distal articulation surface (posteri o rly ) .

2 Lunatum dex. (= Lunare = Os carpi interm e d i u m ) . A = proximal view (anterior side up); B = distal view (anterior side u p ) ; C = lateral view (anterior side to the ri g h t ) . L1 = DAP proximal articulation facet; L2 = DT proximal articulation facet along anterior border; L3 = DT distal articulation facet; L4 = DAP distal articulation facet; L5 = Maximum height (medially ) ; L6 = Maximum DA P ; L7 = Maximum DT.

141 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

APPENDIX 1 (continu e d )

3 Uncinatum sin. (= Uncifo rme = Hamatum = Os carpale quart u m ) . A = proximal view (posterior side up); B = distal view (anterior side up); C = medial view (anterior side to the ri g h t ) . U1 = DAP maximum proximal art i c u l a t i o n f a c e t ; U2 = DT maximum proximal articulation facet; U3 = Maximum DT distal articulation facets (anteri o rly ) ; U4 = M a x i mum height anteri o r ly ; U5 = Maximum height posteri o r ly ; U6 = Maximum DA P ; U7 = Maximum DT.

4 M a g num sin. (= Capitatum = Os carpale tert i u m ) . A = proximal view (anterior side up); B = distal view (anterior side u p ) ; C = lateral view (anterior side to the left). M1 = Greatest diagonal diameter at proximal articulation facet; M2 = Smallest diagonal diameter at proximal articulation facet; M3 = Maximum DAP of distal articulation facet; M4 = Maximum DT of distal articulation facet; M5 = Maximum height; M6 = DAP maximu m ; M7 = DT.

142 MOL et al.: Netherlands proboscideans

APPENDIX 1 (continu e d )

5 Tra p e zoideum sin. (= Os carpale secundum). A = proximal view (anterior side up); B = distal view (anterior side up); C = anterior view.Td1 = Maximum DAP prox i- mal articulation facet;Td2 = DT proximal articulation facet anteri o r ly ; Td3 = Maximum DA P ; Td4 = Height anterior surface m e d i a l ly ; Td5 = Height anterior surface latera l ly ; Td6 = DAP distal articulation facet for Mc. I I ;Td7 = DT distal art i c u l a t i o n facet for Mc II.

6 Trapezium dex. (= Os carpale pri mu m ) . 7 P i s i fo rme dex. (= Os carpi accessori u m ) . A = lateral view; B = posterior view.T1 = Maximu m A = medial view; B = posterior view. P1 = Maximu m L e n g t h ; T2 = DAP maximum proximal articulation facets; l e n g t h ; P2 = DAP maximum prox i m a l ; P3 = DT maximu m T3 = DT maximum proximal articulation facets (posteri o r- p r ox i m a l ; P4 = Maximum DT proximal articulation facet; ly ) ; T4 = DAP maximum distal articulation facet (Mc I);T 5 P5 = Maximum DAP proximal articulation facet; P6 = = DT maximum distal articulation facet;T6 = DT maxi- M i n i mum DAP midshaft; P7 = Minimum DT midshaft. mum midshaft (over tuberosity);T7 = DAP midshaft.

143 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999

APPENDIX 1 (continu e d )

8 Calcaneus sin. (= Os tarsi fi bu l a r e ) . A = proximal view; B = lateral view. C1 = Maximum DT proximal epiphy s i s ; C2 = M a x i mum DT proximal articulation facets; C3 = DAP proximal epiphy s i s ; C4 = DAP proximal articulation facet; C5 = Greatest DAP calcaneum; C6 = DAP tuber calcanei; C7 = Minimum DT tuber calcanei; C8 = Maximum height art i c u l a t i o n facets (in life position); C9 = Maximum height calcaneum (in life position).

9 N aviculare sin. (= Os tarsi centra l e ). A = proximal view (anterior side up); B = anterior view; C = distal view (anteri o r side up). N1 = DAP proximal articulation facet of astra g a l u s ; N2 = DT proximal articulation facet of astra g a l u s ; N3 = DA P m a x i mum proximal articulation facets; N4 = DT maximu m ; N5 = DT maximum distal articulation facets; N6 = DAP distal a r ticulation facets along the centre of the nav i c u l a r e ; N7 = Maximum height nav i c u l a r e ; N7a = Height navicular (anteri o r ly ) m e d i a l ly ; N7b = Height navicular (anteri o r ly) centre; N7c = Height navicular (anteri o rly) latera l ly.

144 MOL et al.: Netherlands proboscideans

APPENDIX 1 (continu e d )

1 0 A s t ragalus dex. A = proximal view; B = distal view; C = posterior view.A1 = DAP maximum proximal art i c u l a t i o n facet (caput tali);A2 = DT maximum proximal articulation facet;A3 = DAP maximu m ; A4 = DT maximu m ; A5 = DA P m a x i mum distal articulation facets (with calcaneus); A6 = DT maximum distal articulation facets (with calcaneus);A7 = M a x i mum height (life position);A8 = Maximum height (with distal articulation facets in hori zontal position).

145 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 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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