Contr. Tert. Quatem. Geol 37(3-1) 39-55 6 figs, 2 pis. Leiden, December 2000
Pliocene species of Aequipecten
(Mollusca, Bivalvia, Pectinidae)
from the North Sea Basin
R. Marquet
Institut royal des Sciences naturelles de Belgique
Brussels, Belgium
and
H.H. Dijkstra
University of Amsterdam
Amsterdam, the Netherlands
Marquet, R. & H.H. Dijkstra. Pliocene species of Aequipecten (Mollusca, Bivalvia, Pectinidae) from the North Sea Basin. — Contr.
Tert. Quatem. Geol., 37(3-4): 39-55,6 figs, 2 pis. Leiden, December 2000.
Pliocene Basin deposits in theNorth Sea have so far yielded three species of the bivalve genus Aequipecten P. Fischer, 1886, viz.A. o.
& opercularis (Linné, 1758), A. r. radians (Nyst Westendorp, 1839)and A. wagenaari n. sp. The last-named taxon appears to be re-
to stricted the Early to Middle Pliocene of the North Sea Basin. In the literature, Aequipecten r. radians has been recorded from a number of Miocene and Pliocene localitiesboth within and the North records beyond Sea Basin; however, these have proved errone- ous. Material from Aquitaine (France) formerly considered to be assignable to A. r. radians shouldbe referred to A. liberatus (Coss- mann & Peyrot, 1914). Material from Touraine and Brittany represents the subspecies, A. r. assimilatus (Millet de la Turtaudière, first in Miocene 1866). Aequipecten opercularis occurs the Lower of Aquitaine, being represented by the subspecies, A. o. pinorum Middle of (Cossmann & Peyrot, 1914). Miocenerecords A. opercularis are known from the Lower Badenianof Romania.
Key words Bivalvia,] Pectinidae, Pliocene, North Sea Basin, taxonomy, new species.
R. Marquet, Institut royal des Sciences naturelles de Belgique, c/o Constitutiestraat 50, B-2060 Antwerpen, Belgium [e-mail: Rob- [email protected]]; H.H. Dijkstra, Department of Malacology, Zoological Museum, University of Amsterdam, P.O. Box 94766, NL-1090 GT Amsterdam, the Netherlands[e-mail: [email protected]].
Contents described species ofpectinid was collected; this had pre- viously been encountered in Dutch beach material, but
not been formally named because of lack of in situ Introduction p. 39 yet material. It is here described Systematic descriptions p. 39 as Aequipecten wagenaari
Conclusions p. 48 n. sp.
Acknowledgements p. 48
References p. 48
SYSTEMATIC DESCRIPTIONS
Introduction Abbreviations — In the text the following abbreviations
are used: IRScNB 1ST - Institut royal des Sciences
Newly collected Neogene molluscan material from har- naturelles de Belgique. Brussels (Belgium); RGM - Na- tionaal Natuurhistorisch Leiden bour, underground and motorway works in the Antwerp Museum/Naturalis. (the Netherlands, area (NW Belgium) has allowed an extensive revision of formerly Rijksmuseum van Geologie en
- bv - bivalved Miocene and Pliocene faunas. In recent years, quite a Mineralogie); RM R. Marquet Colin; numberof systematic studies on Pliocene gastropods and specimen, v - single valve. bivalves have come out (Marquet, 1984, 1993, 1995a, b,
1996, 1997a-c, 1998; Marquet & Dijkstra. 1999). while Order OstreoidaFerussac. 1822
Suborder Pectinina Waller. 1978 iithologic logs and and maps have been published by Pcctinoidea Janssen (1974), Hoedemakers & Marquet (1992) and Superfamily Wilkes. 1810 Pectinidae Wilkes, 1810 Marquet (1998). The present paper continues the series Family
von on Pliocene bivalves from Kallo. At this locality, an un- Subfamily Chlamydinae Teppner. 1922 40
Tribe Aequipectini Waller, 1993 fig. 2 reconditus J. de C. Sowerby, 146, pi. 575, Genus Aequipecten P. Fischer. 1886 1827 Pecten p. figs 5, 6 (non Solander in Brander, 1766).
1843 Pecten opercularis. Lin. — Nyst, p. 291, pi. 23, fig. Type species — Ostrea opercularis Linne, 1758, by 2. monotypv (Recent eastern Atlantic). 1843 Pecten Sowerbyi Nyst, p. 293, pi. 22, fig. 3 New synonyms — Aequipecten (Perapecten) Wagner, 1856 Pecten opercularis Linnaeus — Wood, p. 35, pi. 6, 1985 species, by original designation, is (p. 84), type fig. 2.
commutatus Monterosato. 1875 (= nam. nov. Pecten Pecten J. — pro 1856 gracilis, Sowerby Wood, p. 37, pi. 6, Michelotti. Pecten philippii Recluz, 1853, non 1839); fig. 5.
distribution: Recent, eastern Atlantic, from Portugal 1874 Pecten niveus? Macgillivray — Wood, p. 104, pi.
8. southwards to Senegal and in the Mediterranean). Lin- 8, fig.
1874 Pecten opercularis var. arcuata Wood, p. 105, pi dapecten Petuch, 1995 (p. 40), type species, by original 8, 6. distribu- fig. designation, is Pecten muscosus Wood. 1828; 1878 Pecten opercularis, L. —Nyst, pi. 15, fig. 1. tion: Recent, (sub)tropical western Atlantic. 1878 Pecten sowerbyi, Nyst — Nyst, pi. 15, fig. 2.
Diagnosis — Aequipectini with a modified hinge denti- 1881 Pecten opercularis, Lin. — Nyst, p. 149 {partim). tion and prominent resilial teeth; shape near-orbicular 1937 Chlamys opercularis (Linne 1758)— van Regteren and equilateral; auricles slightly unequal in size. In early Altena, p. 61 (partim).
ontogeny with a granulate micro-ornament, lacking re- 1950a Pecten (Chlamys) opercularis (Linne 1758) —
ticulation or 'herringbone' structure; in early radial Heering, p. 46, pi. 14, figs 3-6 (partim).
1950b Pecten (Linne) — 13, stage, shell generally with ventrally concave commargi- (A.) opercularis Heering, p. 2, 23. nal lirae on rib flanks. pi. fig. 1957 Chlamys (Aequipecten) opercularis Linné, sp. 1758 Remarks — Wagner (1991, p. 42) raised Perapecten to
— 28 Glibert, p. (partim). generic level. However, morphological features of this 1969 Lyropecten (Aequipecten) opercularis (Linnaeus, taxon. i.e. macro- and micro-ornament and auricular 1758) — van Regteren Altena et al., p. 16, pi. 10,
crura, are similar to those of Aequipecten. Petuch (1995, fig. 38; pi. 11, fig. 38 (partim).
considered be different from — p. 40) Lindapecten to 1976 Chlamys (Aequipecten) opercularis (Linnaeus)
the basis of the of more Aequipecten mainly on presence Tebble, p. 60, pi. 5, fig. b, d.
1979 — Lucas, 14. erect spines on radial costae. This, however, is a very Aequipecten opercularis (L.) p. 1981 Chlamys (Aequipecten) 1758) insignificant character, which may also be observed in opercularis (Linne,
— 7. Lauriat-Rage, p. 44, pi. 4, fig. species of the pectinid bivalve genera Chlamys Roding.
1991 — Aequipecten opercularis (L., 1758) Wagner, p 1798 and Mimachlamys Iredale. 1929. The overall mor- 40, text-fig. 22, pi. 6, figs 1-5. phological characters of Lindapecten and Aequipecten 1992 Aequipecten (Aequipecten) opercularis (L., 1758) are similar, which is why they are here synonymised. — Cavallo & Repetto, p. 186, fig. 550. Typical extant species of Aequipecten are the fol- 1993 Chlamys opercularis (Linnaeus, 1758)— Poppe & lowing: Goto, p. 63, pi. 9, fig. 4.
1994 Chlamys (Aequipecten) scabrella (Lamarck, 1819)
- — Ben nonLamarck). A. commutatus (Monterosato, 1875) from the Moussa, p. 83, pi. 3, fig. 12 (
1999 Ostrea Linnaeus, 1758 — Dijkstra, (sub)tropical eastern Atlantic and subtropical West opercularis p fig. 7a-d. Africa; 421,
- from the A. exasperatus (G.B. Sowerby II, 1842)
Locus — France. Caribbean region; typicus Brittany,
Stratum — Recent. - A. flabellum (Gmelin, 1791)from the tropical eastern typicum
Materialstudied — In addition the material Atlantic; to type (see
Dijkstra, 1999, 422), we have studied the following - p. A. muscosus (Wood. 1828) from the (sub)tropical
of A. o. in RM Colin): Ant- western Atlantic; specimens opercularis (all (Borgerhout Ring, Lt. Lippenslaan. 4 bv/3 v); - A. glyptus (Verrill. 1882) from the (sub)tropical werp Rivierenhofand Ring Foorplein, all from western Atlantic; Antwerp (Ring ‘Heterocetus level', Diest Formation, Deurne Member, 1 - A. nicklesi Dijkstra, 1998 from tropical West Africa; bv/12 v); Kallo Lillo Formation, Oor- - A. opercularis (Linne, 1758) from the boreal to sub- (Verrebroekdok, deren Member, Cultellus level, 108 v); Kallo tropical eastern Atlantic; 2 - A. tehuelchus (d'Orbigny. 1847) from the (subtro- (Vrasenedok, upper part Kattendijk Formation, bv); Antwerp (Kruisschanssluis, Lillo Formation, Lucht-bal pical southwestern Atlantic. Member, 4 bv); Antwerp (Noordkasteel. Luchtbal Mem-
ber, 13 bv); Wijnegem (Het Sas, Austruweel Member, 1
v); Ekeren (Schoonbroek, Kruisschans Member, 10 v); Aequipecten opercularis opercularis (Linné, 1758) Kallo (Verrebroekdok, Kruisschans Member, basal level. PI. 1, Figs 1,2 39 v); Kallo (Verrebroekdok, Kruisschans Member, sand
mixed derived * lens, 39 v); Antwerp (Ring Borgerhout. 1758 Ostrea opercularis Linne, p. 698. Pliocene. 1 Haven- 1825 Pecten gracilis J. de C. Sowerby, p. 129, pi. 393, bv); Antwerp (Berendrecht-Negende 41
dok, Merksem Member, 109 v); Kallo (Verrebroekdok, 1914) we have studied 20 valves from Salles (Ruisseau
Merksem Member. 21 v); Gedgrave (The Cliff, Coralline de Dubern, Gironde, Aquitaine, France; Falun de Salles,
Crag, 28 v); Gedgrave (The Broom, Coralline Crag, 21 Middle Miocene, RM Colin).
Sutton Wood. Coralline Sut- — v); (Rockhall Crag, 50 v); Dimensions Height 67.2 nun, length 70.8 mm, di- Park Coralline ton (Sudbourne Pit. Crag. 50 v); Walton- ameter 24.0 mm (bivalved specimen, see PI. 1, Fig. 1); on-the-Naze (The Naze. Essex, Waltonian Red Crag, 11 height 39.2 mm, length 41.7 mm, diameter 12.6 mm (bi-
Alderton Farm Newbournian v); (Buckanay Pit, Suffolk, valved specimen, see PI. 1, Fig. 2). For height and length
Red Crag, 3 v). ofa sample of 50 specimens see Fig. 1.
Of & Aequipecten o. pinorum (Cossmann Peyrot,
Fig. 1. Aequipecteno. opercularis (Linné, 1758),Kallo(Verrebroekdok), Lillo Formation, Oorderen Member, Cultellus level and
basal level (all RM Colln).
Description — Medium-sizedspecies, reaching heights nolium well developed. Hinge line usually straight, of less than 40 mm in the Kattendijk Formation, but up rarely the auricles form an obtuse angle, so that the umbo to 70 mm in the Oorderen Member (Lillo Formation), lies below the auricles. Edge of hinge line turned inside. while extant specimens may grow to heights of 100 mm. Anteriorly, parallel to the hinge line, occurs a plica.
Shell strong, rather tumid, slightly inequilateral and in- Ligament pit shallow and relatively broad. Primary ribs
with left valve flatter than visible equivalve slightly right one. inside, muscle insertion vague.
Ratio of dimensions variable; in Kattendijk Formation, Discussion — The present species is extremely common
while in the Oor- from specimen length may surpass height, at Kallo, and ranges the Kattendijk Formation to deren Member mostly the opposite is true. Disc nearly the Merksem Member (Lillo Formation). Elsewhere in circular, posteriorly slightly oblique. Umbonal angle Belgium, there are Pleistocene (Eemian) beach records. about 105°. Ornament consisting of 18-25 primary radial In Great Britain, A. o. opercularis is recorded from the Red and Mammalian while in the Neth- ribs (20 in fossil material examined, see Fig. 2). which Coralline. crags, erlands it is known from boreholes well are about half as wide as the intercostal spaces. On the as as from mate- central of the disc these ribs three rial washed ashore. part mostly comprise Heering's (1950b. pi. 2, fig. 23) A. secondary ones; near the anterior and posterior borders specimen clearly belongs to o. opercularis, which the difference between primary and secondary disap- means that his material is possibly younger than Mio-
The pears. Intcrcostally very regular, fine growth lines occur, cene. oldest occurrence in Belgium is probably in which into small scales which continue the Late Miocene Deurne Member. Pecten may develop on sowerbyi
for reconditus C. the radial ribs. Auricles unequal, anterior one clearly Nyst, 1843, a nom. nov. Pecten J. de larger than posterior. In the narrow border area between Sowerby, 1825 (non Solanderin Brander, 1766), is syn-
with A. disc and auricles, antimarginal ornament is present, es- onymous o. opercularis, judging from shell pecially anteriorly. Ornament of anterior ear consisting shape and number of radial ribs (22) in Sowerby's fig- of five clear ured radial ribs, posterior auricle with more, but specimens. Nyst (1878) did still illustrate P. sower- but there is mention which finer ribs. Length of both auricles together reaching byi, no in the text appeared in about 50% of total shell length. Byssal notch deep, cte- 1881. 42
2. of the number of radial ribs in from Kallo Lillo Forma- Fig. Histogram Aequipecteno. opercularis (Linné, 1758) (Verrebroekdok),
tion, Oorderen Member, Cultellus level (RM Colln).
* Pecten gracilis J. de C. Sowerby, 1825 is based on 1839 Pecten radians Nyst & Westendorp, p. 15, pi. 3, material from the Red Crag of East Anglia. Differences fig. 11.
— 3b. from 1843 Pecten radians Nyst Nyst, p. 294, pi. 24, fig. A. o. opercularis mentioned by Wood (1856) in-
1856 Pecten dubius, Brocchi — Wood, p. 38, pi. 6, fig. clude a flatter, thinner shell, slightly different size ratios, 3; nonpi. 4, fig. 3 (partim). and smaller auricles. In showing these features, Sow- 1866 Pecten assimilatus - Milletde la Turtaudiere, p. 30 ' erby's form is nearly identical with 'var. lineatus (da 1878 Pecten radians, Nyst — Nyst, pi. 15, fig. 3. Costa. ofA. o. 1778) opercularis. 1881 Pecten — radians, Nyst Nyst, p. 151
to & Goto extant A. o. According Poppe (1993), — 1939 Chlamys radians Nyst Roger, p. 134, pi. 16, figs
in between the Lofoten Is- 8-12 9 =A. opercularis occurs an area (partim, nonpi. 17, fig. r. assimilatus). lands (Norway) in the north and Madeira, the Canary 1945 Pecten (Aequipecten) radians Nyst — Gilbert, p. 5. Islands and the Azores to the south, being also known 65, pi. 3, fig.
1950b Pecten scabrellus Lamarck — Heering, 13, from the Mediterranean, but not from the Black Sea. The (A.) p. 2, 24,28 (non Lamarck). bathvmetric is from low tide level down pi. figs range to 4(H) m. 1957 Chlamys (Aequipecten) radians Nyst, sp. 1839 — Like fossil shells, extant specimens characteristically Glibert, p. 27. have between 18 and 25 radial ribs. 1958 Chlamys scabrella (Lamarck) var. sarmentica Aequipecten o. (Cossmann & Peyrot, 1914) pinorum — Goldfuss Ertlnal-Erentoz, p. 149, pi. 24, fig. 8. from the Sallomacian of (Middle Miocene) Aquitaine 1965 Lyropecten (Aequipecten) radians (Nyst, 1839) — first described distinct Here, (France) was as a species. Glibert& van dePoel, p. 27. we follow Glibert & van de Poel (1965) in considering it 1969 Lyropecten (Aequipecten) opercularis (Linnaeus,
— Altena 16 a subspecies of A. opercularis. Indeed, shell shape and 1758) van Regteren etal, p. (partim). 1986 size radial ribs divided into three Chlamys (Aequipecten) zenonis Cowper-Reed, and primary (each sec-
— 1936 Studencka, p. 40, pi. 4, fig. 11. ondaries, with two intercostal secondary ribs) are all
1995 Aequipecten radians (Nyst, 1839) — Lozano- similar to those of the nominate subspecies, and differ- Francisco & Vera-Pelaez, p. 26. ences are slight. The number of ribs in A. o. pinorum is now 1920 Pecten (Aequipecten) seniensis Lamarck — Dollfus lower and lines form (18-20) commarginal growth & A. Dautzenberg, p. 411, pi. 37, figs 8-20 (= r. as- scales radial thicker when crossing the ornament. This similatus).
with records — subspecies, together ofA. sensu non b opercularis 1970 Chlamys radians Nyst Bongrain, p. 35, pis a, lato from Lapugiu (Romania) cited by Studencka et al. (= A. r. assimilatus).
1970 radians var. assimilis — (1998). represent the first occurrence of the taxon to non Chlamys Nyst (Millet) d A. date. Bongrain, p. 37, pis c, (= r. assimilatus).
non 1981 Chlamys (Aequipecten) radians (Nyst, 1839) —
2 Lauriat-Rage, p. 45, pi. 4, fig. 6; pi. 5, fig. (= A. r. assimilatus). Aequipecten radians radians won 1987 Chlamys radians — Andres Galache & de Porta & (Nyst Westendorp, 1839) 8. Vemet, p. 146, pi. 1, figs 7, PI. 1 2, Fig. won 1994 Chlamys (Aequipecten) radians (Nyst) — Ben 43
Moussa, p. 79, pi. 3, fig. 6. tained in RM Colin) from St. Clement de la Place
(Marne-et-Loire, Le Grand Chauvereau, 'Redonian', 15
Locus typicus — Antwerp (Belgium). v); Sccaux d'Anjou (Marne-et-Loire, La Presselierc.
— Pliocene. Stratum tvpicum Lower •Redonian', 21 v) and Brigne (Beugnon, Marne-et-Loire,
Materialstudied — In addition the Indre-et- to lectotype (IRScNB 'Redonian', 4 v), Ferriere Largon (pit near D66.
1ST 4770), we have studied the following material ofA. Loire, 'Pontilevian', 10 v); Ferriere Largon (Ferme La from r. radians: specimens Edegem (Edegem Member. Placettc, Indre-et-Loire. 'Pontilevian', 59 v); Le Grand
Lower Miocene(?), illustrated by Glibert (1945), Pressigny (Paulmy. Indre-et-Loire. 'Pontile-vian'. 5 v);
IRScNB 1ST 1925, 1926); specimens from Antwerp, Pauvrclay (Carricre Buzele, Indre-et-Loire. 'Ponti-
IRScNB 1ST 'Scaldisien', as illustrated by Nyst (1878), levian', 8 v); Manthelan (quarry near D50, Indre-ct-
4768. 4799, 4800; IRScNB 1ST 6403. here illustrated Loire, 'Pontilevian', 3 v); Le Louroux (Le Petit Bray.
(PI. 2, Fig. 1), as well as the following specimens (all in Indre-et-Loire, 'Pontilevian', 11 v); Amberre (quarry Kat- RM Colin): Antwerp (dock works at Noordkasteel, near Moulin Pocha, Vienne, Touraine. 'Pontilevian', 15 tendijk Formation. Ditrupa level, 15 v); Kallo (Vrasene- v).
Petaloconchus 123 dok, Kattendijk Formation. level, v); Of A. liberatus (Cossmann & Peyrot. 1914) we have
Antwerp (underground Schijn-poort, Kattendijk Forma- studied the following material (all in RM Colin): Mar- tion. Ditrupa level, 4 v); Antwerp (parking Gaumont tillac (Pas de Barreau. Gironde, Burdigalian. Falim de
Cinema. Kattendijk Formation, Ditrupa level, 21 v); Leognan, 22 v); Saucats (Coquilliere, Gironde. Burdiga-
' Antwerp (Borgerhout Ring. Lt Lippenslaan. Heteroce- lian. 27 v); Corbleu (Moulin de Carro. Gironde. Falun de tus level', Diest Formation, Deurne Member, 1 v); and St. Avit, Aquitanian-Burdigalian, 7 v); Saint Martin
Estepona (Velerin Conglomerate, province of Malaga, d'Oney (quarry along D365. Landes. Falun de St. Avit, 2 Spain, Zanclean. v). Aquitanian-Burdigalian. 7 v); Roquefort (Palombieres Of A. assimilatus have studied material from r. we Douze valley, Landes, Falun de St. Avit, Aquitanian-
Ferrière Larson, Pontlevoy and Bossée (Touraine, 'Pon- Burdigalian, 2 v); Uzeste (Moulin de Gamachot. Gi- tüevian'. illustrated as by Dollfus & Dautzenberg, 1920, ronde. Falun de Bazas, Aquitanian-Burdigalian. 25 v).
IRScNB 1ST 1379-1391), as well as material (all con-
Fig. 3. Aequipecten r. radians (Nyst & Westendorp, 1839), Kallo (Vrasenedok), Kattendijk Formation, Petaloconchus
and Similipecten levels (RM Colln).
Dimensions — Height 12.0 mm, length 12.0 mm, di- valves of typical specimens with eighteen very fine, pri-
PI. radial ribs which about half the ameter 4.2 mm (bivalved specimen, see 2, Fig. 1). mary (see Fig. 4), are
For height and length of a sample of 50 specimens see width of intercostal spaces. These spaces show on each
Fig. 3. side of the primary ribs very fine antimarginal lines. In
— lines these Description Small species, up to 23 mm in height, addition, clear growth occur; become irregu- with strongly inequivalve, but only slightly inequilateral, lar where they cross secondary radials. On the primary
shell. of formed of radial fragile Shape disc near ventral margin circular, ribs, spines are at junction and commar- but part of anterior and posterior margin closest to lire ginal ornament, on the posterior side occasionally scales umbo straight insteadof curved (from about half height). occur. Many specimens have in the centre ofthe disc 4-5
Both valves strongly flattened. Ornament variable and radial ribs which are heavier than the other primary
different with often markedly on left and right valves. Right commarginal ones, better-developed spines. 44
Fig. 4. Histogram of the number of radial ribs in Aequipecten r. radians (Nyst & Westendorp, 1839),Kallo (Vrasenedok), Kattendijk Formation, Petaloconchus level (RM Colln).
Left valve invariably with finer ornament, usually lack- cian' strata in Aquitaine and in 'Pontilevian' deposits of
ing spines or heavier commarginal growth lines. In some Touraine, is characterised by a higher number of primary
A. number of specimens, the ornament of the right valve also is mark- ribs composed of, as in opercularis, a sec-
edly weaker, while all radials arc equally well devel- ondary ones. These ribs are as wide as the intercostal
radial ribs, scales rather than oped. Usually the secondary radials and a few spines spaces; on the spines are remain visible. Hinge line straight, auricles large, taking present, and specimens with heavier primary ribs are
the shell sizes than up about half of total shell length, posterior auricle al- absent. Moreover, may grow to larger
ways clearly smaller than anterior one. Where disc and that ofA. radians. The general shape, however, is nearly auricles meet, antimarginal ornament occurs. Auricles the same as in A. radians, being also flattened In other
with 5-7 (anterior) or 3-5 (posterior) radial ribs, about as features, however, A. liberatus is closer to A. opercu-
wide as interspaces. Inner side of shell with small, ob- laris.
tuse triangular ligament pit and a clearly marked ridge, Another representative of tire same group occurs in
parallel to hinge line. Ribs visible interiorly only up to the Rcdonian" of Brittany and in the Pontilcvian' of
half height. Ctenolium internally clearly visible, with 4-5 Touraine. Dollfus & Dautzenberg (1920) wrongly identi-
of this material ‘Pecten denticles. Muscle insertion nearly indistinguishable. fied part as (Aequipecten) seni-
— Poel Discussion The present species is abundant through- ensis Link', while Roger (1939), Glibert & van de
out the Kattendijk Formation at Kallo and at other lo- (1965) and Lauriat-Rage (1981) used the specific name
calities around Antwerp; is not rare either in the Pliocene radians for it. Differences include the higher number of
(Coralline Crag) of Great Britain and in Dutch material. radial ribs (c. 20), narrower intercostal spaces, much
There are a few records of Miocene material from the finer commarginal ornament, mostly lacking spines or
North Sea Basin; Glibert (1945) listed specimens from scales on radials, and a more tumid shell. However,
- ribs the Early Miocene Houthalen and Edegem members specimens with stronger radial do occur; see e.g.
these constitute Dollfus & 37, 20, IRScN 1ST appear to the earliest record of the spe- Dautzenberg (1920, pi. fig.
' cies, which is also found in the Late Miocene Hetero- 1391) and Bongrain (1970, pis c, d). In addition, ac-
Sands' the rib number cetus (Diest Formation). Heering (1950b) re- cording to Roger (1939, p. 135), primary
corded, under the name of Pecten (A.) scabrellus, mate- may amount to 18. For that reason, this material is here
rial from Maasbree (Limburg, the Netherlands), which, considered to be closely related to the North Sea Basin judging from his illustrations, should be referred to A. r. species, but assignable to a distinct subspecies which is radians. restricted to the Atlantic 'Redonian' and 'Pontilevian-
Previous authors who followed Roger (1939) consid- Savignean' (Miocene). For this type of shell, the name ered two Miocene species to be synonymous with A. Pecten assimilatus Millet de la Turtaudiere, 1866 is
radians, although they are not. Aequipecten liberatus, available. Ben Moussa (1994) identified specimens of
which in and 'Salloma- Messinian from A. radi- occurs Aquitanian. Burdigalian (Late Miocene) age Morocco as 45
studied the in RM ans; the material he illustrated shows only 13 radial ribs we have following specimens (all
different that of A. Kallo Oorderen An- and a shell shape completely from Colin): (Verrebroekdok. Member, radians. Studencka's (1986, pi. 4. fig. 11, under the gulus benedeni level, 2 v); Kallo (Verrebroekdok. Oor-
of from deren Atrina 1 Kallo name Chlamys (Aequipecten) zenonis) specimen Member. level. v); (Verrebroekdok.
Oorderen Member, basal level. 14 Kallo Rybnica (Poland), of Middle Miocene age. represents v); (Verre- Cultellus 66 genuine A. r. radians with its typical ornament. The broekdok. Oorderen Member, level, v);
same probably applies to a Middle Miocene specimen Antwerp (Ring Borgerhout, mixed derived Pliocene, 3
from the Karaman Basin (Turkey) figured by Eriinal- v); Orford (The Broom. Suffolk, Coralline Crag. 2 v);
Erentoz (1958. pi. 24, fig. 8) with typical radians orna- Sutton (Sudbourne Park Pit. Suffolk. Coralline Crag. 1
ment and shell shape. Kojumdgieva & Strachimirov v); Walton-on-the-Naze (The Naze, Essex, Waltonian
(1960, pi. 22, figs 9-12) illustrated material from the Red Crag, 2 v).
the of Of A. malvinae Bois de Middle Mioceneof Bulgaria, under name Chlamys (du Montpereux. 1831). we
(Aequipecten) seniensis (Lamarck), which certainly does have studied one valve from Lychow (Tarnobrzeg. Po- of Middle Miocene not belong to that species, on account of their symmetri- land) age (RM Colin).
cal shell shape. Their shape in fact is rather like that of OfA. elegans (Andr/.cjowski. 1820). we have studied
A. radians, but illustrations of the ornament are not clear material from Weglin (Tr/ydniczanca valley, Tanobrzeg,
Poland. Middle Miocene. 19 RM A. enough to be certain of this; material from the region v, Colin), of vasco- was not available for study. Studencka's, Eriinal- niensis (Cossmann & Pevrot, 1914), specimens from
Erentoz's and possibly also Kojumdgieva & Strachimi- Mios (Ruisseau de Surgenne. Gironde. Falun de Mios. of rov "s records are younger than the oldest North Sea Ba- 'Sallomacian', 6 v, RM Colin), A. degrangei (Coss-
mann & valves from Salles (Ruis- sin records of A. r. radians, which thus appears to have Peyrot, 1914), eight Falun de Middle Mio- migrated from the North Sea Basin into the Paratethvs. seau de Minoy, Gironde. Salles,
Apparently, Aequipecten radians also reached the Atlan- cene. RM Colin), of A. raouli (Cossmann & Peyrot. and level tic "Pontilevian' and 'Redonian' localities, where a dis- 1914), two valves from the same locality (RM
tinct subspecies developed, probably because of separa- Colin), and of A. seniensis (Lamarck. 1819) material Berchem tion from the main distribution area. In the Mediterra- from Antwerp (Ploegstraat 29, Formation,
or records to Member, level, 21 RM nean, no authentic Miocene Recent appear Antwerp Panopea v, Colin): 25 RM exist, while Pliocene specimens are known only from the Antwerp (Schijnpoort. same level, v, Colin);
Iberian Peninsula. Baldichieri (orchard on road to Monale, Asti, Italy, Mid-
radians RM and Diolo Aequipecten cannot be considered a prede- dle Pliocene, 23 v, Colin), (calanca on
cessor of A. o. opercularis, which rather appears to have road to Chiavenna, province of Piacenza. Italy. Piaccn-
Miocene of Middle 37 and 1 RM evolved from A. o. pinorum from the Middle zian. Pliocene. v bv. Colin).
the Aquitaine Basin. Dimensions — Height (holotype) 18.6 mm. length 18.4
mm, diameter 4.6 mm; paratvpe IRScNB 1ST 6402 (left
valve), height 19.3 mm. length 19.5 mm. diameter: 4.0
Aequipecten wagenaari n. sp. mm; paratype RGM 456 050 (right valve), height 20.7
PI. 2, Fig. 2 mm. length 20.0 mm, diameter 5.6 mm. For height and
length of a sample of 50 specimens see Fig. 5.
1856 Pecten — — dubius, Brocchi Wood, p. 38, pi. 4, fig. 3 Diagnosis Rather small (c. 30 mm in height) species
(parti m). with near-circular, strongly tumid disc, with a primary
1881 Pecten opercularis, Lin. — Nyst, 149 (partim). p. radial rib number of 15. without secondary radial orna-
1937 Chlamys opercularis (Linne 1758) — van Regteren ment, with rather heavy commarginal ornament, forming 61 Altena, p. (partim). scales on junctions with radial ornament, and with auri- 1950a Pecten (Chlamys) opercularis (Linne 1758) — cles which together measure about 3/4 to 2/3 of total Heering, p. 46, pi. 14, figs 7-9, 25 (partim). shell 1738 length. 1957 Chlamys (Aequipecten) opercularis Linné, sp.
— Rather small with a maximum — 28 Description species, Gilbert, p. (partim).
of 32.0 mm shell solid, in- 1969 ? — height Fig. 5); slightly Lyropecten (Aequipecten) spec van Regteren (see
16. Altena et al., p. equilateral and inequivalve. Disc nearly orbicular, but height mostly exceeding length. Shell rather tumid, left
Derivatio nominis — In honour of Mr M. Wagenaar, for valve flatterthan right. Ornament at first consisting of 15
his valuable help in producing the photographs which primary radial ribs (see Fig. 6). which in the majority of
illustrate this and previous papers. specimens are not divided into secondary ribs; this oc-
Locus — Kallo of typicus (Verrebroekdok), province curs only exceptionally in the largest specimens. Width Oost-Vlaanderen, Belgium. of radial ribs about a third of the intercostal spaces on
Stratum — Formation, Oorderen Member third the tvpicum Lillo left valve, to a on right valve, which is usually Middle Pliocene. (Cultellus level). more strongly ornamented than the left one. Growth
— is IRScNB 1ST form low scales in intercostal Types Holotvpe 6401, paratvpes are lines spaces, on the ribs
IRScNB 1ST 6402 and RGM 456 050. are high, hollow scales. Hinge line nearly always
Material studied — In addition to the type specimens. straight. 46
Fig. 5. Aequipecten wagenaarin. sp., Kallo(Verrebroekdok), LilloFormation, Oorderen Member, Cultellus level and basal level (RM
Colln).
Fig. 6. Histogram of the number of radial ribs in Aequipecten wagenaari n. sp., Kallo (Verrebroekdok), Lillo Formation, Oorderen Member, Cultelluslevel and basal level (RM Colln).
Auricles comparatively large, to more than 3/4 of total Discussion — The present species differs from A. oper-
length in juvenile specimens and 2/3 in adults. Anterior cularis in having a smaller shell (see Figs 1 and 5), in auricle twice as large as posterior, with a deep byssal shell length exceeding height, in usually having about
ribs of notch. In the area between disc and auricles occurs anti- fifteen radial instead twenty, in nearly always
marginal ornament, especially anteriorly. Anterior auri- having single radial ribs instead of divided into secon-
cle with six radial ribs, posterior one with three. Anteri- dary ribs, in showing on the radial ribs heavy scales, in
Auricular line, and in orly, clear scales are present on the radials. having nearly always a straight hinge having
denticles anteriorly clearly visible, ctenolium well devel- relatively more tumid valves with wider auricles.
oped, with five teeth visible on the inside of the shell. Differences between the new species and A. r. radi-
Ligament pit broadly triangular, shallow. Hinge line with ans also are clear. Aequipecten wagenaari has a much ridges, parallel to margin. Radial ribs clearly visible on more tumid shell, the ornament on left and right valve thicker the inside ofthe shell, muscle insertion vague. differs less, radials are and their number is lower 47
of has (15 instead of 18), intercostal secondary radial lines are The Miocene western Paratethys yielded, ac- which invariably absent, and the radials have scales instead of cording to Studencka et al. (1998), eight species
to the A. spines; moreover, the umbonal portions of anterior and possibly belong genus Aequipecten: opercu-
much limited laris, A. A. radians and A. seniensis posterior margin are straight to a more macrotis, r. (=
degree. scabrellus), all of which have been discussed above, as A. Bois In Miocene strata of Belgium, a single species of well as A. elegans, A. malvinae, diaphanus (du
Aequipecten occurs, namely A. seniensis. This was well de Montpereux, 1831) and A. lapugyensis (Nicorici, malvinae differs in described and illustrated by Glibert (1945, pp. 69, 72, pi. 1977). Aequipecten clearly having a Stu- 3. fig. 6; pi. 4, fig. 2). It differs even more clearly from much higher number of radial ribs (29-40; see dencka. 4, 7, Material of.A. A. wagenaari in shell shape (i.e. moderately to strongly 1986, p. 35, pi. figs 5, 10). inequilateral and oval instead of near-circular); its rib elegans contained in the RM Colin has only 10-13 radial
of the which but which be di- number is mostly comparable to that new species, ribs, are mostly smooth, may ribs. The but the radials are relatively wider and consist of at least vided into fine secondary commarginal orna-
ribs. Scales is weak and while two, but often more secondary are prominent ment very occurs only intercostallv,
scales on the radial ribs are absent. Its is between as well as on the radial ornament elements. spines or shape
in the Miocene of oval rather than circular, it A. Aequipecten seniensis occurs so more closely resembles.
Touraine. although some of the records of this species seniensis. Aequipecten diaphanus, figured by Friedberg
in fact Dollfus & and 147, are erroneous identifications by (1936, p. 230, pi. 39, figs 1-3) Roger (1939, p.
Dautzenberg (1920). It is also known from the pi. 18, fig. 11) differs markedly from A. wagenaari in the shell Paratethvs; the youngest records are from the Pliocene of having radial ornament near margins only,
Italy and Spain. By that time, however, the species had while the centreof the disc remains smooth. Miocene of the gone extinct in the North Sea Basin. In strata Aquitaine occur a large Another Miocene species from Belgium has also number of species ofAequipecten. Unlike A. wagenaari,
vasconiensis Cossmann & 123. been previously considered to belong to the genus A. (see Peyrot, 1914, p. Middle Aequipecten, viz. Mimachlamys angelonii (Meneghini, pi. 14. figs 22-25; pi. 16, figs. 1-6) from the has with 18 1859). This differs from A. wagenaari in having 20-25 Miocene an oval, strongly inequilateral shell, in the central radial ribs, bearing 3-5 rows of strong spines; scales are wide radial ribs, rounded in cross section;
intercostal of the disc these radials of never present and the commarginal ornament part are not composed secon- the anterior and is relatively weak. The shells grow to much larger sizes, daries. They are, however, near posterior
in The in A. vasconiensis reaching almost 70 mm height. Mimachlamys angelo- margins. commarginal ornament
nii is known from the Italian and Austrian Miocene, is highly variable; in some specimens it is nearly absent,
from where it has been described under the name of in others developed into clear scales on the ribs and in-
Chlamys glaesneri Kautsky, 1928. Studencka et al. tercostally. This might represent but a forma of sympa-
(1998) assigned it to the genus Amussiopecten Sacco, tric A. seniensis, which has the same shell shape, but a
1897. differentornament.
and level is A. The Middle-Late Miocene A. r. assimilatus of Tou- Another species from the same area
raine and Brittany differs from A. wagenaari in having degrangei (see Cossmann & Peyrot, 1914, p. 132. pi. 20. is and much flatter than A. more radial ribs (c. 18), finer commarginal ornament figs 5-8), which larger and radial without scales or spines and relatively wider auricles. wagenaari, has wider auricles more numerous
The tumid, near-circular and nearly equilateral shell ribs (25-28); secondary ornament is absent; commarginal
and scales are on and between the radials. shape, however, is quite similar and these species may lines present
be closely related. Aequipecten liberatus has more (20) but are weak. This species is not unlike A. r. assimilatus,
but the shell is flatter and lias and wider ribs than A. wagenaari, and these are com- grows larger; markedly & posed of secondary ribs. Its shell is flattened and grows more radial ribs. Aequipecten raouli (see Cossmann
to twice the size ofthat ofA. wagenaari. Peyrot, 1914, p. 124, pi. 16 [not 14 as indicated in the
Aequipecten macrotis (G.B. Sowerby, 1847) was text], figs 40-42) lias flat valves, very wide auricles, 18-
originally described from Miocene (Tortonian) strata 20 radial ribs, wider than intercostal spaces, is near-
near Lisbon (Portugal). Material from the type area was equilateral but height exceeds length; the commarginal
ornament is like that of the from illustrated by Roger (1939, p. 114, pi. 14, figs. 7-10; pi. preceding species,
15. figs 7, 8; text-fig. 58), who considered Chlamys which this taxon mainly differs in shell shape and in
(Aequipecten) multiscabrella Sacco, 1897, Ch. plubien- number of ribs. Aequipecten raulini (see Cossmann &
sis Ugolini, 1906, Ch. (A.) vasconiensis, Ch. (A.) raulini, Peyrot. 1914, p. 128, pi. 17. figs 1. 2, 22. 23), which also
Ch. (A.) raouli, Ch. (A.) degrangei and Ch. (A.) liberata occurs in Middle Miocene deposits, is near-equilateral,
With the of with wide auricles and rib number the to be subjective junior synonyms. exception a high (24-25); with the second, these taxa are here considered to be valid. commarginal ornament is heavy, prominent scales
Portuguese material of A. macrotis differs from A. intercostally as well as on the radials; it is much smaller
wagenaari in having 18-24 radial ribs with small squa- than A. wagenaari. haveri 1847 also considered mae, but without spines, whileits disc shape comes close Pecten Michelotti, was
to that ofA. opercularis. by Cossmann & Peyrot (1914, p. 134, pi. 17, figs 6-9) to 48
in the belong to Aequipecten; however, this is incorrect. Both deren Member (Lillo Formation) Antwerp area and the and be also in Dutch borehole and valves are too inequilateral ornament resembles appears to common
it is A. that of Chlamys spinulosus (von Minister in Goldfuss, beach material. However, never as common as o.
1833) more closely than it does that of any representative opercularis, and would seem to constitute a species en-
of.Aequipecten. demic to the Pliocene of the North Sea Basin.
Sacco (1897) discussed a lot of Neogene species
which he considered to be assignable to Aequipecten. malvi- Amongst these are A. opercularis, ‘A.’ haveri, A. CONCLUSIONS
nae, and A. seniensis; all of these have been discussed
above. In the Miocene of Torino A. occurs northamptoni Species of Aequipecten are difficult to identify on ac- this is than three times (Michelotti, 1839); more larger count of a wide range of variation in ornament, which
than A. wagenaari, and has. judging from Sacco's (1897. often differs between both valves. Therefore, no single
pi. 4, figs 1-16; pi. 5, figs 1-11) figures fifteen radial ribs feature may be considered diagnostic for any of its spe-
(18-20 according to the text on p. 16), which are most cies. but a set of characters in a population rather than in
of several secondaries and bear clear often composed a single individual shell are needed to define taxa. As the
spines in typical specimens. Aequipecten bicknelli Sacco, histograms of Figs 2, 4 and 6 show, the number of ribs 1897 is found in the Piemonte (p. 21, pi. 6, figs 14-17) however forms a Gauss curve in the three sympatric spe-
Pliocene; it has 25-28 strongly spinose. narrow radial cies of Aequipecten from the Pliocene of the North Sea
ribs, and its shell is rather flattened and round, being Basin. A summation of the three frequency distributions
than twice the size of that of A. with clear more wagenaari. would never give a normal curve a primary 1897 Aequipecten spinovatus Sacco, (p. 21, pi. 6, figs value. Thus this character appears to be of real impor-
21-25). from the Pliocene of Italy, is close in shell shape tance. In addition, variability in itself could be consid-
bicknelli and also has number radial ribs to A. a large of ered a character. This is notably the case in A. radians,
These little wider than those of the of with (22-25). are a previ- where the occurrence specimens stronger pri-
centre ous species and show in the of the disc a tendency mary ornament is a featureof populations of this species,
to split into three secondaries. even if not every individual shows it. When this feature
Of the MioceneA. mioalternans 1897 Sacco. (p. 23, is altogether absent, the population can never be consid- these pi. 7, figs 11, 12) only fragments are known; are ered to represent A. radians. In other forms under study.
with 16-17 primary radial ribs and secondary ones A. more features re- large, e.g. liberatus, differentiating might
in the intercostal ornament is veal themselves careful examination. spaces. Commarginal upon multiscabrellus 1897 29, from lacking. Aequipecten Sacco, (p. The genus Aequipecten appears to have spread
pi. 8, figs 38-41) from Miocene strata in Italy is higher different centres of speciation in the European Neogene.
than long, tumid, with 19-22 radials and long auricles; The Miocene North Sea Basin, from which A. radians
and the ribs centrally, commarginal ornament spines on seems to have migrated into the Paratethvs and the At-
the anterior and are absent, but near posterior margins lantic and the Aquitaine (France), whence the A. oper-
these be The species is not unlike the into the may present. cularis stock appears to originate, spreading Miocene A. raulini. scabriuscu- Aquitaine Aequipecten North Sea Basin in the latest Miocene. In addition, sev-
lus (Matheron. 1842). from the Miocene of the Rhone eral endemic species evolved in the Atlantic, Mediterra-
and from illustrated Sacco Valley (France) Italy was by nean and Paratethys basins during the Miocene and Plio-
30. 9. it is the A. (1897, p. pi. figs 1-4); a very large species, cene, youngest of which was wagenaari, which
with 14-17 radial ribs composed of secondaries; the shell was confined to the Middle Pliocene of the North Sea
is widerthan high. TheEarly Miocene ‘A.’ oblitaquensis Basin.
Sacco. 1897 (p. 18. pi. 5, figs 12-14) probably belongs to
another genus, on account of the shape ofits left valve.
Van Altena the first mention Rcgteren (1937) was to ACKNOWLEDGEMENTS
the occurrence of A. wagenaari in Dutch beach material,
but did name it. recorded not Heering (1950a) specimens For allowing us to study material contained in the
' of Pecten from boreholes sunk (Chlamys) opercularis’ IRScNB and RGM collections, and to use IRScNB re-
at Beers and Oss with sixteen ribs, all of Vlissingen, search facilities, we wish to thank AV. Dhondt. A.W.
the he which probably belong to new species. However, Janssen, F. Wesselingh and K. Wouters. M. Wagenaar, considered these the of to belong to var. sowerbyi Nyst L. Cilis (IRScNB) and W. Miseur (IRScNB) assisted
A. As outlined this was based opercularis. above, variety with the photography, for which we are grateful.
on specimens with 22 radial ribs. Aequipecten wagenaari also Wood's occurs in the British Pliocene, since (1856,
4, indeed has 16 the shell pi. fig. 3) specimen ribs; shape REFERENCES
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PLATE 1
1. bivalved Kallo Lillo Fig. Aequipecten o. opercularis (Linné, 1758), specimen (RM Colln), (Verrebroekdok), Formation, Oorderen
Member, Atrina level); A - right valve, x 1.1; B - left valve, x 1.1; C - both valves, x 1.4.
Fig. 2. Aequipecten o. opercularis (Linné, 1758), bivalved specimen (RM Colln), Kallo (Vrasenedok), Kattendijk Formation, top
level; micro-ornament, A - x 2.5; B - x 6.8. 53\1 54
PLATE 2
1. radians bivalved IST Kallo Fig. Aequipecten r. (Nyst & Westendorp, 1839), specimen (IRScNB 6403), (Vrasenedok), Kattendijk
B - left C - Petaloconchus - micro-ornament of left Formation, level); A right valve, x 3.6; valve, x 3.7; valve, x 11.4 (SEM
images).
Fig. 2. Aequipecten wagenaari n. sp., Kallo (Verrebroekdok), Lillo Formation, Oorderen Member, Cultellus level; A-C - holotype
left and - left (IRScNB IST 6401), valve, x 2.5, x 2.4, x 5.6, respectively; D paratype (IRScNB IST 6402), valve, x 3.8. 55\2
PLATE 2