October 2003 CainozoicResearch, 2(l-2)(2002), pp. 135-137,

Mactromeris polynyma (Stimpson, 1860)

(, )

from the Upper Pliocene of the Netherlands and Belgium

Peter+W. Moerdijk

Kingstraat 14, NL-4336 LG Middelburg; e-mail: [email protected]

Received: 10 March 2003; revised version accepted 16 April 2003

in The Arctic surfclam, Mactromeris polynyma, is recorded from strata of Late Pliocene (Reuverian) age the North Sea Basin for the

first time; this represents a Pacific invader which became extinct in , but survived to the present day along the eastern

American coast.

KEY WORDS:: Mollusca, , Mactridae, Pliocene, North Sea Basin.

Introduction predominantly of Pliocene and Pleistocene age, a search

was conducted for in situ records of M. polynyma. Two

A hinge fragment of a large fossil mactroid, dredged from complete valves are now known from the Late Pliocene

the Westerschelde (the Netherlands), which could not be (Reuverian) Kruisschans Member(and possibly Merksem

attributed to any of the known species from the southern Member; LilloFormation) in the Antwerp area (Belgium).

North Sea Basin, is here assigned to the extant Arctic surf These had previously been attributed to arcuata

Subse- The illustrations , Mactromeris polynyma (Stimpson, 1860). (Sowerby, 1817). present paper provides

quently, more identifiablefragments have been recognised ofM. polynyma from the North Sea BasinNeogene; differ-

in fossil collections from the Westerschelde. In view ofthe ences between this species and coeval mactroids in this

fact that dredged fossil shells from the Westerschelde are area are discussed.

Figures 1, 2. Mactromeris polynyma (Stimpson, 1860),ROM 456 091, from Antwerp (Belgium), 9th harbour dock. Pliocene, Lillo

Formation, Kruisschans Member (or Merksem Member), length of valve 105 mm.

Description and distribution ures 1, 2) is large, up to in excess of c. 130 mm, with the

umbo slightly prosogyrate.The anterior end is lower than

The near-equilateral shell of Mactromerispolynyma (Fig- the posterior one and narrowly rounded, somewhat cu- - 136-

neate.The anterior dorsal margin is either straight or con- from typical M. polynyma. Unfortunately, Wood did not cave; broadly rounded at the posterior end, more or less illustrate, nor describe, the valve’s interior; his material has truncated.The ventral is curved. The margin hinge shows a not yet been retraced. It cannot be ruled out that this shell

large resilifer, forming a bulge in the ventral margin ofthe belongs to M. polynyma as well.

The hinge plate. lateral teeth are short and unomamented Specimens ofS. arcuata from the Pliocene of Iceland,

in The sinus is illustrated (not grooved as Spisula). pallial quite deep by Gladenkov et al. (1980), are more equilateral and confluent with the line. than venfrally not pallial typical S. arcuata and show a more strongly curved

At present, M. polynyma occurs in the boreal Pacific ventral margin. In fact, they are more closely similar to and northwest Atlantic, having been recorded from the Sea Wood’s S. procrassa than to S. arcuata. Judging from the of Okhotsk, Bering Sea, Alaska, south to Puget Sound and quite long lateral hinge teeth they obviously cannot be from Hudson Bay to Massachusetts, living in mud of assigned to M. polynyma.

from low in al. exposed areas the intertidal zone to depths of Coan et (2000) presented an extensive synonymy of about 110 metres (Coan et al, 2000). From the fossil M. polynyma and were of the opinion that a more careful record, Marincovich (1983) mentionedthis species from as examination might well lead to the recognition of one or early as the Middle Miocene of Alaska, Japan and eastern more subspecies. The handful of specimens and fragments

Siberia. The aims to documentthe from the North Sea Basin be shell present paper presence cannot distinguished on of M. polynyma in the Netherlands and Belgium during the outline, characters of the hinge and pallial/adductor scars

Late Pliocene (Reuverian, according to the standard used from extant M. polynyma.

The earliest record from by TNO-NITG, Utrecht). eastern Mactromerispolynyma is a representative of a faunal

North America is thatof Richards (1962), who listed it as invasion from the Pacific into the northern Atlantic which

fossil from the Pleistoceneof Maine. a occurred during the Middle and Late Pliocene and Early

Pleistocene (Meijer, 1993), being one of the cooler-water

ofthis fauna. with Yoldia components Together e.g. myalis Discussion (Couthouy, 1838), Megayoldia thraciaeformis (Storer,

1838) (see Janse et al., 2003) and Mya arenaria Linne,

shell from the Westerschelde and the 1758, M. of Pacific bivalve Previously, fragments polynyma represents a group two specimens from the Pliocene of Antwerp have been immigrants which became extinct in Europe but survived confused with Both & the the Spisula arcuata. Geys Marquet to present day along eastern American coast. The

(1979, pi. 35, fig. 4; pi. 36, fig. 1) and Janssen et al. (1984, absence of this species from deposits both older and

illustrated valves of under fig. 135a, b) M. polynyma the younger than the Late PlioceneKruisschans Member (and

of S. That is well known in the name arcuata. species possibly Merksem Member) is a reflection of a relatively

North Sea Basin Pliocene illustrations short of (for good see Wood, presence M. polynyma in the North Sea Basin. It is

1857, pi. 23, fig. 5; Nyst, 1878, pi. 24, fig la-c; Marquet, of note that the oldest record from eastern North America

1993, 2, The lateral pi. figs 3,4). quite long, grooved hinge is of Pleistocene age. teeth of S. arcuata clearly demonstrate that it is a repre-

sentative ofthe - genus Spisula. In addition, Spisula arcuata Materialstudied RGM 456 091 (leg. P.A.M. Gaemers, differs from M. polynyma in having a less equilateral and 13 March 1976, 1 left valve, Antwerp, 9th harbour dock; smaller shell (up to c. 82 mm), and in having a convex Pliocene, Lillo Formation, Kruisschans Member (or Merk- anterior dorsal Other mactroid margin. large species de- sem Member); 1 right valve (present whereabouts un- scribed from the Pliocene of the North Sea Basin are: known), Antwerp, Kallo harbour works, Lillo Formation,

Mactra glauca Bom, 1778, M. stultorum Linne, 1758, Kruisschans Member (see Geys & Marquet, 1979, pi. 35,

and Spisula artopta (Wood, 1857) S. procrassa (Wood, fig. 4; pi. 36, fig. 1); RGM unregistered (leg. de Vreede, 1

1857). Of these, M glauca mostly closely resembles M. fragment ofright valve (+ fragments of right 7 valve and polynyma, but its anterior dorsal margin is convex and the fragments of shell edge?), Westerschelde, near Ellewouts- ventral margin of the hinge plate is almost straight; in dijk (province of Zeeland, the Netherlands), derived from addition, it has longer lateral hinge teeth than M. polynyma. Pliocene sediments; RGM 395 748 (leg. H. Smits, 1950-

Mactra stultorum is decidedly smaller than M. polynyma, 1954, 2 fragments of left valve, Westerschelde near Elle- its anterior dorsal margin is also convex rather than straight woutsdijk, derived from Pliocene sediments); 1 fragment of or concave and the lateral hinge teeth are longer. Spisula right valve (A. Goetheer Colin, 23 August 1997), Wester- is related S. from the artopta clearly to arcuata judging schelde near Baarland (province of Zeeland, the Nether- inequilateral shell, but is triangular in outline and more lands), derived from Pliocene sediments. inflated. Both Spisula artopta and S. procrassa have grooved lateral hinge teeth.

Wood (1879) mentioneda single specimen of ‘Mactra Acknowledgements ponderosa Stimpson’, albeit with a query, from the Red

Crag of Waldringfield, sent to him by a Dr Reed, who had Many thanks are due to F.P. Wesselingh (Nationaal identifiedthe shell after as M. ponderosa Stimpson (1851). Natuurhistorisch Museum, Leiden) for providing literature,

Mactra ponderosa Philippi, 1844 (non Eichwald, 1830, making materialavailable and commenting on earlier drafts

is M. nec Conrad, 1830) a junior of polynyma. of this paper, to G.J. Vermeij (Department of Geology,

Judging from the illustration, however, this shell differs University ofCalifornia at Davis) for reviewing the manu- - 137-

script and providing valuable information on the fossil leontology 84, 59-155. Marquet, R. 1993. The molluscan fauna of the record of the present species, to T. Meijer (TNO-NITG, Kruisschans Member (Lillo Formation, Late Pliocene) in the Antwerp area Utrecht) for commenting on an earlier typescript and (Belgium). Contributions to Tertiary and Quaternary Geol- making available material for examination, R. Marquet 83-103. ogy 30, (Antwerp) and A. Goetheer (Ovezande) for making mate- Meijer, T. 1993. Stratigraphical notes on Macoma (Bivalvia) in rial available for study, and to G.A. Peeters (Schiedam) for the southern part of the North Sea Basin and some remarks permission to use his excellent drawings. on the arrival ofPacific species. In: Janssen, A.W. & Janssen,

R. (eds). Proceedings of the Symposium ‘Molluscan Palae-

ontology’, 11th International Malacological Congress, Siena, References Italy, 30th August-5th September 1992. Scripta Geologica,

Special Issue 2,297-312.

Bom, I. von 1778. Index rerum naturalium Museum Caesarei Nyst, P.H. 1878. Conchyliologie des terrains tertiaires de la 458 Vindobonensis 1, Testacea, xlii + pp. Wien (Kraus). Belgique, I. Terrain pliocene Scaldisien. Annales du Musee Coan, E.V.,Valentich Scott, P. & Bernard, F.R. 2000. Bivalve d’Histoirenaturelle de Belgique (Paleontologie) 3 (atlas), 28 of western North America. Marinebivalve mollusks pis. from Arctic Canada to Baja California. Santa Barbara Mu- Philippi, R.A. 1844. Descriptiones testaceorum quorundam seum ofNatural History Monograph2 [Studies of Biodiver- novorum, maxime chinensium. Zeitschriftfur Malakozoolo- sity, 2], 764 pp. gie 1(11), 161-167. T.A. 1830. On the Conrad, geology and organic remains of a part Richards, H.G. 1962. Studies on the marine Pleistocene, part II: of the peninsula of Maryland. Journal of the Academy of The marinePleistocene mollusks of eastern North America. naturalSciences ofPhiladelphia 6, 205-230,pis 9, 10. Transactions theAmerican of philosophical Society, n.s. 52, Naturhistorische Eichwald, E. von 1830. Skizze von Lithauen, 42-141. Volhynien und Podolien in geognostisch-mineralogischer, Sowerby, J. 1817. The mineralconchology ofGreat Britain; or 256 3 botanischer und zoologischer Hinsicht, pp., pis. Wilna. colouredfiguresand descriptions of those remains oftesta- Geys, J.F. & Marquet, R. 1979. Veldatlas van de Cenozoische ceous or shells, which have been preserved at vari- fossielen van Belgie. Publikatievan de Belgische Vereniging and ous times depths in the earth 2(28), 129-140, pis 157- 125 voor Paleontologie2, pp. 162. London (Meredith). Gladenkov, Yu. B., Norton, P. & Spaink, G. 1980. [Upper Stimpson, W. 1851. Shells of New England. A revision of the Cenozoic of Iceland (Stratigraphy of Pliocene-Pleistocene New with synonymy of the testaceous mollusks of England, and Paleontological assemblages)]. Academy ofSciences of notes their and their on structure, geographicaldistribution, the USSR, Transactions 345, 1-116 (in Russian) with 57 Boston figures ofnew species, pp. (Phillips, Sampson Janse, A.C., Moerdijk, P.W. & Meijer, T, 2003. First record of & Co.). Megayoldia thraciaeformis (Storer, 1838) (Bivalvia) from the Stimpson, W. 1860. Check lists of the shells ofNorth America, Pleistocene of the North Sea Basin. Cainozoic Research 2, 3. East coast: Arctic seas to Georgia. Smithsonian miscella- 137-142. neous Collections2(128), 1-6. der Janssen, A.W., Peelers, G.A. & van Slik, L. 1984. De fossiele S.V. 1851-1857. A Wood, monograph of the Crag mollusca, or, schelpen van de Nederlandse stranden en zeegaten, tweede description of shells from the Middleand Upper Tertiaries of serie, 8 (slot). Basteria 48, 91-219. the east of England, 2. Bivalves. Monograph of the Palae- C. Linne, von 1758. Systema naturae per regna tria naturae, ontographicalSociety, London, 341 pp. secundum classes, ordines, genera, species, cum characteri- Wood, S.V. 1879. Second supplement to the monograph of the bus, differentiis, synonymis, locis. Editio decima reformata Middle Crag mollusca, or, description of shells from the and 824 1(1-2), iii+ pp. Holmiae (Laurenti Salvii). Upper Tertiaries of the east of England. Monograph of the Marincovich, L. 1983. Molluscan paleontology, paleoecology, 58 Palaeontographical Society, London, pp. and North Pacific correlations of the Miocene Tachilni For-

mation, Alaska Peninsula,Alaska. BulletinsofAmerican Pa-