on mastodon remains from the netherlands: an ove r … · 55 francien i.b r a b e r 1,dick mol1...

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55 Francien I. Braber 1 , Dick Mol 1 & John de Vos 2 1 Natuurmuseum Rotterdam 2 Naturalis, Leiden On mastodon remains from the Netherlands: an overview Braber, 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.) - ELEPHANTS HAVE A SNORKEL! P APERS IN HONOUR OF P AUL Y. SONDAAR - 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 arvernensis (CROIZET ET JOBERT, 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. arvernensis and Mammut borsoni (HAYS, 1834) found in the Netherlands is given. Correspondence: Francien Braber and Dick Mol, Natuurmuseum Rotterdam, P.O.Box 23452, NL-3001 KL Rotterdam, The Netherlands. John de Vos, Naturalis - National Museum of Natural History, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands. Keywords: Anancus, Mammut, Plio-Pleistocene, Early Pleistocene, Oosterschelde, The Netherlands INTRODUCTION Anancus arvernensis (CROIZET ET JOBERT, 1828) had a wide Eurasian geographical range and existed from the Miocene until the Early Pleistocene. Its largest expanse was during the Pliocene. The oldest known fossils of Anancus arvernensis were found in Miocene deposits in Spain. A l a rge amount of fossils of Anancus was found in Bulgaria in the region of Dorkovo in 1985, in a Pliocene deposit (Thomas et al. 1986). These dental fossils are characterised by a rather primitive morpholo- gy, which either indicates an Early Pliocene form or an intraspecific variation of the spe- cies. Other localities where Anancus arvernen - sis was found are in France (Chilhac), Italy (Valdarno) and the Netherlands (Maalbeek, Liessel, Schelde estuary). In France and in the Schelde estuary the remains of Anancus arv e r - nensis were found in association with Mammuthus meridionalis. Late representatives of Anancus are characterised by molars with cementum in the valleys between the cusps. Both localities in France and the Netherlands show this identical late evolutionary stage of Anancus teeth, which results in a comparable dating of the fossils to 1,9 million years old (Moraal 1998, Boeuf 1993). This means that the Anancus fossils found in the Oosterschelde belonged to the stratigraphically youngest representatives of the species. Anancus became extinct in the Early Pleistocene. Anancus arvernensis had a bunodont dentition and had two tusks pointing straight forward (Fig. 1). The shoulder height of the animal was about 2 m. The mandibles were short and the molars had nipple-shaped cusps arranged in transverse rows. These bunodont molars suggest a soft diet of leaves, fruit and twigs. The feet were made to walk on soft soil, which means that Anancus was an inhabitant of the moist woodlands. In the fossil record the molar crown became higher, the valleys between the cusps and rows were filled with cement and the transverse rows or lophids increased in number. The trend developed from soft forest food towards material like grasses, requiring grinding teeth (De Vos et a l . 1998).

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55

Francien I. B r a b e r1, Dick Mol1 & John de Vo s2

1 N a t u u r museum Rotterd a m2 N a t u r a l i s , L e i d e n

On mastodon remains from the Netherlands:an ove rv 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 claypit near the village of Maalbeek, near Tegelen, The Netherlands. Also, an overview of the known molarsof 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-3001K 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-Pleistocene, Early Pleistocene, Oosterschelde, The Netherlands

I N T RO D U C T I O NAnancus arvernensis (CR O I Z E T E T JO B E RT,1828) had a wide Eurasian geographical rangeand existed from the Miocene until the EarlyPleistocene. Its largest expanse was during thePliocene. The oldest known fossils of A n a n c u sa rv e r n e n s i s were found in Miocene depositsin Spain. A l a rge amount of fossils of A n a n c u swas found in Bulgaria in the region ofDorkovo in 1985, in a Pliocene deposit(Thomas et al. 1986). These dental fossils arecharacterised by a rather primitive morpholo-g y, which either indicates an Early Plioceneform or an intraspecific variation of the spe-cies. Other localities where Anancus arv e r n e n -s i s was found are in France (Chilhac), Italy( Valdarno) and the Netherlands (Maalbeek,Liessel, Schelde estuary). In France and in theSchelde estuary the remains of Anancus arv e r -n e n s i s were found in association withMammuthus meridionalis. Late representativesof A n a n c u s are characterised by molars withcementum in the valleys between the cusps.Both localities in France and the Netherlandsshow this identical late evolutionary stage of

A n a n c u s teeth, which results in a comparabledating of the fossils to 1,9 million years old(Moraal 1998, Boeuf 1993). This means thatthe A n a n c u s fossils found in the Oosterscheldebelonged to the stratigraphically youngestrepresentatives of the species. A n a n c u sbecame extinct in the Early Pleistocene.Anancus arv e r n e n s i s had a bunodont dentitionand had two tusks pointing straight forward(Fig. 1). The shoulder height of the animalwas about 2 m. The mandibles were short andthe molars had nipple-shaped cusps arrangedin transverse rows. These bunodont molarssuggest a soft diet of leaves, fruit and twigs.The feet were made to walk on soft soil,which means that A n a n c u s was an inhabitantof the moist woodlands. In the fossil recordthe molar crown became higher, the valleysbetween the cusps and rows were filled withcement and the transverse rows or lophidsincreased in number. The trend developedfrom soft forest food towards material likegrasses, requiring grinding teeth (De Vos et a l. 1998).

R E S U LT SN ew material of Anancus arve r n e n s i sAnancus arv e r n e n s i s is known from the pro-vince of Zeeland, in the south-western part ofthe Netherlands. Detailed descriptions andfigures of dental remains of Anancus arv e r -n e n s i s are given by Schreuder (1944, 1945)and by Hooijer (1953, 1991). Van Essen &Mol (1996) and De Vos et al. (1998) gave anoverview on the Early Pleistocene mammal-ian remains from the Oosterschelde in theZeeland province. Most of the knownremains of Anancus arv e r n e n s i s are kept inNaturalis, the National Museum of NaturalHistory in Leiden (see Appendix). One molarof Anancus arv e r n e n s i s belongs to the Brabercollection (Delft, The Netherlands) and bearsthe number B0100. This molar was unpublish-ed so far. The exact locality where thisA n a n c u s molar was found is not known, butit was dredged from the bottom of the sea bya vessel from a fisherman from the island ofSchouwen Duiveland. Bryozoa are clearlyrecognisable on the surface of the molar.Since there are other finds of A n a n c u s k n o w nfrom the Oosterschelde which borders the

island, this is the most probable locality. T h emolar became part of the private collection ofmed.dr J.L. Braber in the sixties, when it wasgiven to him as a token of appreciation formedical treatment.

The fossils from the Oosterschelde demon-strate a typical state of fossilisation: whentapped on with a hard object they produce ahigh-pitched sound. The molar in the Brabercollection also has those characteristics and isvery well preserved (Table 1, Fig. 2). Only atthe right front side a cusp has been broken.The colour of the fossil is black for the ena-mel and dark brown for the cementum.Between the cusps there are relatively heavycementum deposits. The molar is slightlyworn at the conules of the protocone and atthe conelets of the paracone, the protoconeitself is missing (broken). The other cones,conelets and conules are unworn. The rootsof the tooth are missing. The molar is an M3dex. with 6 lophs of two cones, of which the6t h loph is much smaller than the other five.In the first loph both cones are accompaniedby two conelets (buccal side of the sulcus) or

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ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

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

two conules (lingual side of the sulcus). Inthe 2n d, 3r d and 4t h loph there is only one cone-let or conule. In the two last lophs each conehas two conelets and two conules, except forthe smallest cone, which has only one cone-let. The cones and conules do not form astraight ridge, but do alternate each other in away that the lingual cone is situated more tothe front than the buccal cone of the samel o p h .

The Anancus arve r n e n s i s molar fro mM a a l b e e kA famous site of Early Pleistocene faunalremains is Tegelen in the province ofL i m b u rg, The Netherlands. Several clay pitsnear Tegelen have yielded large quantities offossils. Most of them originate from the samedeposits (Kortenbout van der Sluijs &Zagwijn 1962). The main fauna of Tegelen isknown from the stage Tiglian C 5, often writ-ten as TC5, and dated with an age of approxi-mately 1.6 million years. From a site calledthe Van Cleef Pit or Maalbeek Pit near thevillage of Maalbeek in the neighbourhood of

Tegelen, much older mammals such asTa p i rus arv e r n e n s i s and Anancus arv e r n e n s i sare known (We s t e r h o ff et al. 1998). Only oneset of Ta p i ru s molars (Kortenbout van derSluijs 1960) and one molar of A n a n c u s(ST96438) were found. Often we comeacross both species mentioned in the faunallist of Tegelen. There has been a lot of dis-cussion on the age of the Maalbeek Pit( We s t e r h o ff et al. 1998). According to manyresearchers the clay of the Maalbeek Pit isplaced in the Eburonian (e.g. Zagwijn 1963).

The molar of A n a n c u s was collected by themuseum staff of the Leiden GeologicalMuseum on February 23, 1960. G. Kortenbout van der Sluijs, under theleadership of professor I. M. van der Vlerk, atthat time was doing a research project on theTegelen fauna. Kortenbout van der Sluijswrote on that day in his Tegelen diary (whichis now in the collection of Dick Mol, Fig. 3):

‘Met Prof.v.d. Vlerk eerst naar Russel-Tiglia. We krijgen hier een aantal beende-ren, o.a. een stuk gewei van E u c l a d o c e ro sen een snijtand van een paard. Smeets is echter ziek, zodat een deel van de vondsten ontvindbaar is. Vervolgens naar Maalbeek. 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 sand 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 molarwas found in situ (1998: 61). This is not quite

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BRABER et al.: mastodons from the Netherlands

Ta ble 1 Measurements in cm of the Anancus arv e rn e n s i s

molar B0100 (coll. B ra b e r ) .The enamel thickness, measured at

the side where the protocone has been broke n , v a ri e s

b e t ween 0.53 cm and 0.81 cm.

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ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

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 ]

a

b

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consistent with the notes Kortenbout van derSluijs wrote in his diary (see above) on hisproject on the Tegelen fauna. The workers in thepit donated the molar to him. We s t e r h o ff et al.(1998) gave some interesting information (p. 62):

‘The new exposures in 1993 and 1994, additional palynological research and a reappraisal of the primary data have

resulted in alternative ideas about the stratigraphical age of the fauna from Maalbeek. The clay with the mastodont molar is here referred to the Tiglian B pollen zone. The tapir remains might be slightly older and can probably be assigned to the Tiglian A pollen zone. The new results indicate that the mammal fauna

BRABER et al.: mastodons from the Netherlands

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

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ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

mentioned above is not younger but older than the fauna from the Tegelen pits.’

This means that Kortenbout van der Sluijs,when he wrote his notes in his Tegelen diary,was correct in his assumption about thedating of a Lower Tiglian age of theMaalbeek Pit. The molar (ST96438) nowbelongs to the collection of Naturalis inLeiden. It is the only A n a n c u s fossil everfound in Maalbeek. This is the first time it isdescribed, measured and depicted (Table 2,Fig. 4). ST96438 is an upper M1 dex., whichis very worn down. On the lingual side of themolar all cusps are totally worn. On the buc-cal side five lophs can be distinguished, allworn.

An ove rv i ew of the Anancus arve r n e n s i smolars found in the NetherlandsThe Appendix provides an overview of all themolars and molar fragments known from theNetherlands. 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 showcomparatively heavy cementum depositsbetween the cusps and therefore belong to thelate evolutionary stage. The localities werethe A n a n c u s and the M a m m u t molars werefound in the Netherlands and in the NorthSea are indicated on the map (Fig. 5).

An ove rv i ew of the M a m mut bors o n imolars found in the NetherlandsMammut borsoni ( HAY S, 1834) is a zygodontmastodon known from the Pliocene of Europeand Northern Asia. The molars have sharptransversal ridges, derived from two fusedcones, specialised for cutting the food likescissors. There are only two or three molarsof Mammut borsoni found in the Netherlands.Two were found at the locality Liessel, a gravel pit in the province of North Brabant;Li 1001 is in the collection of the MuseumJan Vriends in Asten and the other one in thecollection of mr. Nies in Deurne. They werefound at about 100 meters distance from eacho t h e r. The third molar (NMR 698), which is

in the collection of the NatuurmuseumRotterdam, is of unknown provenance. MolarLi 1001 was found in 1998 at a depth of 20m e t e r. It is a crown of a left m3 with five ridges, of which the first four are called lophids and the last the talonid. It is heavilyworn, especially at the buccal side, whichindicates that it belonged to an old animal.The second molar from Liessel was found in1990 at a depth of 22 meters. It is the com-plete crown and a part of the root of a left m3(Peters et al. 1991).

The stratigraphy of the Liessel site is verycomplicated, because the sediment was dredged from the pit. The fossils derivedfrom the pit are diverse; remains of marinemammals dated into the Miocene and fossilsof terrestrial mammals of Pliocene age. A l s oplant remains from the Pliocene were reco-vered. A Pliocene age for the molars ofMammut borsoni is most convenient, like theother terrestrial mammals and the plant seeds.

Ta ble 2 Measurements in cm of the Anancus arv e rn e n s i s

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

t h i c k n e s s , measured on the occlusal of the molar, v a ri e s

b e t ween 0.29 cm and 0.65 cm.

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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 ) .

BRABER et al.: mastodons from the Netherlands

a

b

c

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ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

The third specimen (NMR 698) is only a part(two ridges) of a lower m3. There is no loca-lity information available. Although Vo nKoenigswald (1950) interpreted it as a fossilfound in the Oosterschelde, this is improba-ble. There are no traces of sea-life on the sur-face. The fossils found in the sea always havebarnacles or bryozoans stuck to the surface.Another problem is that Mammut borsoni i sknown from the Pliocene and the finds of theOosterschelde are younger, at least of EarlyPleistocene age.

Mammut borsoni is often found associatedwith Anancus arv e r n e n s i s. They must havehad a different diet according to their denti-tion; Mammut borsoni cutting branches andtwigs with its zygodont teeth, Anancus arv e r -n e n s i s with its bunodont teeth browsing onleaves, both in the moist woodlands at theriver banks.

Figure 5 Sites (underlined names) of all the known molars of Anancus arv e rn 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]

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BRABER et al.: mastodons from the Netherlands

C O N C L U S I O N SThe only A n a n c u s molar which is knownfrom Maalbeek is an upper M1 dex. Its geo-logical age is Tiglian B. Kortenbout van derSluijs (1960) was the first author to recognisethat the Maalbeek fauna was older than themain fauna of the famous Tegelen clay pit.All the published molars and molar fragmentsof Anancus arv e r n e n s i s known from theNetherlands are listed in this paper and all themolars show the heavy cementum depositsbetween the valleys of the cusps. This indica-tes that they all had the same evolutionarystage and therefore we can conclude thatAnancus arv e r n e n s i s became extinct in theEarly Pleistocene in North-western Europe,probably during the Tiglian B. Mammut b o r s o n i is often found associated withAnancus arv e r n e n s i s, but only restricted tothe Pliocene, while A n a n c u s is also knownfrom the Early Pleistocene (Mol & Van Essen1996). In Dorkovo (Thomas et al. 1986) thetwo proboscideans were also found associa-ted, they are of Pliocene age and the larg emolars of Anancus arv e r n e n s i s represent themore primitive stage. In Liessel, where thestratigraphy is rather complicated and sedi-ments could be mixed, M a m m u t and A n a n c u sare both present, but it is not clear that theyderive from the same deposit. BecauseA n a n c u s represents the late evolutionarystage in dentition, characteristic for the EarlyPleistocene age, and the age of M a m m u tfossils is restricted to the Pliocene in Europe,it is assumed that the layers of Late Plioceneand Early Pleistocene age in Liessel aremixed up and that the remains of the twoProboscideans derived from different layers.

AC K N OW L E D G E M E N T SThe authors would like to express their gratitude and admiration to dr Paul Sondaarfor his continuous and ongoing enthusiasm todevelop the Pleistocene vertebrate palaeonto-logy both on a national and global scale withgreat skill. We would like to thank dr JelleReumer for his valuable comments on thisp a p e r, Ko Sturkop for the reconstruction of

Anancus arv e r n e n s i s and Jaap van Leeuwenfor drawing the map of the Netherlands.

R E F E R E N C E SBoeuf, O., 1993 - Il était une fois… Il y a près de

2.000.000 d’années à Chilhac, Haute-Loire, France - Amis de Chilhac (Association)

De Vos, J., Mol, D. & Reumer, J.W. F., 1998 - Early Pleistocene mammalian remains from the Oosterschelde or Eastern Scheldt (province of Zeeland, The Netherlands) - Mededelingen Nederlands Instituut voor Toegepaste Geowetenschappen TNO 60: 173-185

H o o i j e r, D.A., 1953 - On dredged specimens of A n a n c u s,A rc h i d i s k o d o n and E q u u s from the Schelde Estuary, The Netherlands - Leidsche Geologische Mededelingen 19: 255-256

H o o i j e r, D.A., 1991 - Fossielen vissen voor de weten-schap; veertig jaar Kor en Bot (1951-1991) - Kroniek van het Land van de Zeemeermin (Schouwen Duiveland): 1-16

Kortenbout van der Sluijs, G. & Zagwijn, W.H., 1962 - An introduction to the stratigraphy and geology of the Tegelen clay-pits - Mededelingen Geologische Stichting NS 15: 31-37

Kortenbout van der Sluijs, G., 1960 - The fossil tapir of Maalbeek, Netherlands - Publicaties Natuurhistorisch Genootschap in Limburg 12: 12-18

Mol, D.J. & van Essen, J.A., 1990 - Mammut borsonifrom the Netherlands - Lutra 33 (2): 183-186

Moraal, J.M., 1998 - Genootschapsverzamelingen, de mastodont van A u v e rgne - Zeeland, Tijdschrift van het Koninklijk Zeeuws Genootschap der We t e n -schappen 7 (2): 69-74

Peters, A., Lammers, Th. & Mol, D., 1991 - Mastodonten-kiezen uit Liessel (Noord-Brabant) - Cranium 8 (2): 89-96

S c h r e u d e r, A., 1944 - Upper-Pliocene Proboscidea out of the Scheldt and the Lower Rhine - Leidsche Geologische Mededelingen 14: 40-58

S c h r e u d e r, A., 1945 - De mastodont en de olifant onder de “zwarte fossielen” uit de Zeeuwsche wateren - Verhandelingen van het Geologisch en Mijnbouwkundig Genootschap 14: 437-448

Thomas, H., Spassov, N., Kodjumdgieva, E., Poidevin, J.L., Popov, V., Sen, S., Ta s s y, P. & Visset, D., 1986 - Résultats préliminaires de la première mission

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paléontologique franco-bulgare à Dorkovo (arrondis-sement de Pazardjik) Bulgarie - Comptes rendus de l’Académie des Sciences Paris 302, II (16): 1037-1 0 4 2

Van Essen, H. & Mol, D., 1996 - Plio-Pleistocene proboscideans from the southern bight of the North Sea and the Oosterschelde, The Netherlands - in: Shoshani, J. & Ta s s y, P. (eds.) - The Proboscidea, Evolution and Palaeoecology of Elephants and their relatives - Oxford University Press: 214-224

Von Koenigswald, G.H.R., 1950 - Voorlopige medede-ling omtrent het voorkomen van Mastodon borsoni in

Nederland - Geologie en Mijnbouw 12 (1): 14-15We s t e r h o ff, W.E., Cleveringa, P., Meijer, T., Van

Kolfschoten, M. & Zagwijn, W.A., 1998 - The Lower Pleistocene fluvial (clay) deposits in the Maalbeek pitnear Tegelen, The Netherlands - Mededelingen Nederlands Instituut voor Toegepaste Geowetenschappen TNO 60: 35-65

Zagwijn, W.H., 1963 - Pollen-analytic investigations in the Tiglian of the Netherlands - Mededelingen van de Geologische Stichting NS 16: 49-69

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BRABER et al.: mastodons from the Netherlands

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

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DEINSEA - ANNUAL OF THE NATURAL HISTORY MUSEUM ROTTERDAMP. 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