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The occurrence of the genus Fusulculus Bouchet & Vermeij (, Pseudolividae) in the Eocene of England, with...

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The occurrence of the genus Fusulculus Bouchet & Vermeij (Gastropoda, Pseudolividae) in the Eocene of England, with a description of two new species

JEAN-MICHEL PACAUD & STEVE TRACEY

Abstract: Re-examination of Early Eocene shells from Lower Swanwick, Hampshire, previously considered to be juvenile Pseudoliva laudunensis (Defrance), has shown them to represent a new species. Similarly, older Early Eocene shells from Crondall, Hampshire, formerly thought to be Turridae and recorded as Oenopota prisca (Deshayes), represent a second new species. Both species are now considered to be Pseudolividae and are referred to Fusulculus Bouchet & Vermeij, a genus of small shells with a biconical profile and relatively high spiral sulcus, recently recorded in the Palaeogene of England for the first time. They are described as Fusulculus alienopriscus and Fusulculus curryi spp. nov. Fusulculus perminutus comb. nov. is proposed for Buchozia perminuta Cossmann from the Marnes de Gan.

Resumé: Un nouvel examen des coquilles de l'Eocène inférieur de Lower Swanwick , Hampshire, précédemment rapportées à des exemplaires juvéniles de Pseudoliva laudunensis (Defrance), montre qu'il s'agit d'une espèce nouvelle. D'autres exemplaires, plus anciens de l'Eocène inférieur de Crondall (Hampshire), et qui furent rapportés par erreur au Turridae Oneopota prisca (Deshayes), correspondent à une seconde espèce nouvelle. Ces deux espèces sont considérées ici comme des Pseudolividae et sont rapprochées de Fusulculus Bouchet & Vermeij, un genre de petite taille au profil biconique et au sulcus spiral relativement haut, récemment signalé dans le Paléogène anglais. Elles sont décrites sous les noms de F. alienopriscus nov. sp.et de F. curryi nov. sp. Fusulculus perminutus (Cossmann) nov. comb est aussi proposé pour Buchozia perminuta Cossmann qui provient de l'Eocène des Marnes de Gan.

J.-M. PACAUD, Laboratoire de Paléontologie, Muséum National d’Histoire Naturelle, 8 rue Buffon – F–75005 Paris. S. TRACEY, University of Greenwich, Environmental and Earth Sciences, Pembroke, Chatham Maritime, Kent ME4 4TB accepted:: 14th February 2000

INTRODUCTION the Dennis Curry collection by the Natural History Museum, The shell figured as Pseudoliva laudunensis (Defrance, London, specimens from the latter, older horizon have been 1826) from the Early Eocene London Clay Formation of examined and found to represent another undescribed Lower Swanwick, Hampshire (see Pl. 1, fig.1) by Jeffery & species of Fusulculus. These specimens become the type Tracey (1997) is closely similar to a number of examples series of Fusulculus curryi sp. nov. collected from different Hampshire Basin localities (London Clay Formation, division B: Kingsclere and Nursling, Abbreviations Hampshire; division A3: Southleigh, and others). All of The specimens examined are now in the following these individuals are of small size and have the appearance repositories: of juveniles. However, comparison with immature examples BMNH - Palaeontology Department of the Natural History of P. laudunensis from the Lower Eocene of Pourcy (Marne) Museum, London. and Sinceny (Aisne) in the Paris Basin (Pl. 1, fig. 5) shows this to be a quite different species. The true P. laudunensis MNHN – Laboratoire de Paléontologie, Muséum National does in fact occur in the English Early Eocene; adult shells d’Histoire Naturelle, Paris. are known from both the Harwich Formation, Oldhaven Member at Swanscombe, Kent and Aveley, Essex and from SYSTEMATIC PALAEONTOLOGY the London Clay Formation at Kingsclere, Hampshire and Bognor Regis, West Sussex (Tracey, 1992; Pacaud, 1998). Class GASTROPODA Cuvier, 1797 The two species are therefore approximately coeval, and Order APOGASTROPODA Salvini-Plawen & while the new species may prove to be rather common, Haszprunar, 1987 actual juvenile examples of P. laudunensis have not yet been Suborder Cox, 1960 recorded from English deposits. These facts have previously Section Thiele, 1929 tended to obscure the separate identities of the two species. As in the case of various other Palaeogene species listed Superfamily MURICOIDEA da Costa, 1776 below, this small new species displays all the teleoconch Family PSEUDOLIVIDAE Gregorio, 1880 characters of the recently-described genus Fusulculus Bouchet & Vermeij, 1998, based on two living deep-water Genus Fusulculus Bouchet & Vermeij, 1998 species from the south-west Pacific. The shell from Lower Swanwick figured by Jeffery & Tracey (1997: pl. 10, fig. 23- Type species by original designation: Fusulculus crenatus 24), is here selected as the holotype of Fusulculus Bouchet & Vermeij, 1998, Recent, New Hebrides Arc, alienopriscus sp. nov. south-west Pacific. Specimens recorded as Oenopota prisca (Deshayes) were Remarks: Until now, the only pseudolivid genus recorded listed by King & Curry (1992) in the faunas of both the from England has been Pseudoliva Swainson, 1840. London Clay Formation and also the Tilehurst Member English Early to Middle Eocene sediments have yielded the (now part of the Harwich Formation) of Crondall and Up relatively large and globose species P. fissurata (Deshayes, Nately, Hampshire. Since the recent acquisition of part of 54 PACAUD & TRACEY

1835), P. laudunensis (Defrance, 1826) [=Buccinum England semicostatum Deshayes, 1835 non Brocchi, 1814], and P. F. pusillus (Beyrich, 1854) [Purpura] Germany & England obtusa (Deshayes, 1835), all since attributed by Vermeij (1998) to Sulcobuccinum d’Orbigny, 1850. However, only two pseudolivid species, “Pseudoliva” nodulosa (Beyrich) and “P.” pusilla (Beyrich), are recorded from the overlying Fusulculus alienopriscus sp. nov. Middle Eocene Selsey Formation (Tracey et al., 1996) and Plate 1, figs.1, 2 Barton Clay Formation (Wrigley, 1941). These two differ from the former group of species in being much smaller, somewhat juvenile in appearance, with a fusiform shape and v 1992 Oenopota (Buchozia) prisca (Desh.); King & Curry: 143, 145 (non Deshayes, 1862) (in part: relatively high spire, a subsutural ramp on the last whorl, London Clay Formation sensu stricto references axial ornament beginning at the suture and gradually only). becoming obsolete below the subsutural ramp, and an obscure spiral sulcus that is shallow and wide, and situated a v 1997 Pseudoliva laudunensis (Defrance); Jeffery & short distance below the periphery. These are the characters Tracey: 97, pl.10, figs. 23, 24 (non Defrance, of the recently described pseudolivid genus Fusulculus 1826) Bouchet & Vermeij, 1998, which has since been recognised v 1998 Fusulculus sp. Pacaud: 12. as being a distinct taxon as early as the Maastrichtian in the v 1999 Fusulculus sp. Pacaud & Schnetler: 62 southeastern U.S.A. (Pacaud & Schnetler, 1999). The two Derivatio nominis: From the Latin alienus, foreign, and new Early Eocene species described here, F. alienopriscus priscus in view of its similarity to the French species and F. curryi, show teleoconch characters similar to those of Fusulculus priscus (Deshayes, 1862). the two living species, Fusulculus crenatus and F. albus Bouchet & Vermeij, 1998, and can be referred to this genus Holotype: BMNH PI TG 1111 (S. Tracey coll.). with confidence. Locus typicus: London Clay Formation, division Cl, the The continuity of the fossil record of Fusulculus from the former Bursledon Brickworks at Lower Swanwick, Late Cretaceous through the Palaeogene is illustrated by the Hampshire. following summary of known species (after Pacaud, 1998; Paratypes: BMNH PI TG 10480 - 10492, (D. Curry coll.), Pacaud & Schnetler, 1999): BMNH GG 22545 (A. Wrigley coll.), GG 22546 (J. Cooper Maastrichtian: & D. J. Ward coll.), GG 22547 (A. Scott coll.), GG 22549 F. macnairyensis (Wade, 1917) [Conorbis] Tennessee, U. (S. Tracey coll.), division C, Fair Oak nr. Kingsclere, S.A. Hampshire, 19 examples in all. Danian: BMNH GG 22548 (C. King & S. Tracey coll.), division C, F. multinodulosus (Vermeij, 1998) [Sulcobuccinum] Nursling nr. Southampton, Hampshire, 4 examples; GG Belgium. 22551, 1 juvenile example (S. Tracey coll.). Selandian: BMNH GG 22550 (S. Tracey & A. Lawson coll.), division F. koeneni (Ravn, 1939) [Pseudoliva] Denmark. A3, Southleigh nr. Havant, Hampshire, 5 examples. F. nanapullus (Pacaud & Schnetler, 1999) Greenland. Other material: J-M. Pacaud coll., lot no. P46299, division Thanetian: A3, Southleigh nr. Havant, Hampshire, 2 examples. F. rosenkrantzi (Traub, 1979) [Pseudoliva] Austria. Distribution: Early Eocene (early Ypresian) of the F. priscus (Deshayes, 1862) [Etallonia] Paris Basin. Hampshire Basin, England; known from the London Clay F. antiquus (Vincent, 1878) [Pseudoliva] Wansin, Belgium. Formation, divisions A3 to C1. early Lutetian: Diagnosis: A small Fusulculus with relatively low-spired, F. perminutus (Cossmann, 1923) [Buchozia] comb. nov. broadly biconical shell, ornamented with thick axial ribs in Gan, France. mid-whorl, forming a nodular shoulder, and with spiral cords mid Lutetian to Priabonian on base. Teleoconch whorls separated by impressed sutures, F. nodulosus (Beyrich, 1854) [Purpura] Germany & bordered by a subsutural collar (bourrelet). Spiral sulcus shallow. Protoconch multispiral.

specimen height diameter aperture length Aperture width

Holotype PI TG 1111 L.Swanwick 3.7 2.4 2.2 0.9 Paratypes GG 22549 a Kingsclere 2.9 1.8 1.9 0.9 b Kingsclere 2.9 1.7 1.6 0.9 c Kingsclere 2.5 1.5 2.0 0.8 Paratypes GG 22548 a Nursling 2.6 1.5 1.6 0.9 b Nursling 2.4 1.4 1.6 0.8

Table 1. Measurements (mm).of Fusulculus alienopriscus sp. nov. EOCENE GASTROPOD 55

Measurements (mm): see table 1. subsutural ramp, absent in both F. nodulosus and F. pusillus. Unlike those two species, F. alienopriscus bears a Description: Shell thick, solid, ovoid, consisting of 3 spiral ornament which is restricted to the abapical part of protoconch and c.3 teleoconch whorls. Larval shell the whorl below the spiral sulcus, and is visible only on the naticoid, of 3 smooth whorls with a thin, almost orthocline last whorl. F. alienopriscus is closer to certain Palaeocene lip (Pl. 1, fig. 2). Teleoconch whorls convex with a species such as F. koeneni (Ravn, 1939) from the Selandian distinctly concave subsutural ramp occupying nearly half of of Copenhagen, F. priscus (Deshayes, 1862) from the Paris the exposed height of the spire whorls. Axial ornament Basin and F. curryi sp. nov. from the Tilehurst Member of consisting of 10-15 low, rounded costae per whorl, the Harwich Formation. These species are all of somewhat truncated below by the spiral sulcus above by the shoulder, similar shape, tapering towards the base. However F. where they form indistinct tubercles. The costae are more or alienopriscus differs in having a more broadly conical last less orthocline on the last whorl, but on the spire they are whorl, and coarse spiral ornament limited to the base, below very variable, and may also form strongly opisthocyrt a very shallow spiral sulcus. F. nanapullus Pacaud & angles or curves on any one whorl. In fresh examples the Schnetler, 1999 from the Palaeocene of Nuussuaq (West shell surface is glossy with a subnacreous sheen, especially Greenland), differs from F. alienopriscus by the absence of in mid-whorl, whereas less fresh shells show dense, slightly a true subsutural ramp on the last whorl. sinuous, imbricated collabral laminae. Spiral ornament consists of a thick subsutural bourrelet adapically, and 6 The multispiral protoconch of this new species suggests that cords below the spiral sulcus, the most adapical one nodular a planktotrophic larva is plesiomorphic for the genus. The at its intersection with the growth lines, the most abapical paucispiral protoconch of Recent Fusulculus species, one very indistinct. Spiral sulcus shallow, terminating in a indicating non-planktotrophic development, is considered to triangular labial tooth. Aperture ovate with broadly open be the derived condition as the evolution of non- siphonal notch. Columella simple, lip not thickened, planktotrophic from planktotrophic larvae has occurred regularly convex except for a slight constriction at level of independently in many other gastropod families. In view of subsutural ramp and of sulcus. the close similarity of its teleoconch morphology to that of the Recent type species Fusulculus crenatus Bouchet & Discussion: This new species is not an immature stage of Vermeij, 1998, F. alienopriscus can confidently be referred Pseudoliva laudunensis (Defrance). The difference is to this genus. shown well by comparison of similarly sized examples of juvenile P. laudunensis (Pl. 1, fig. 5) and adult F. alienopriscus (Pl. 1, fig. 1). When young, P. laudunensis Fusulculus curryi sp. nov. has a regularly pyriform shape, similar to that of the adult shell, with poorly developed sinuous ribs, and at no stage of Plate 1, fig. 3 its ontogeny shows an ovate-fusiform profile, subsutural ramp nor any nodular ornament on the shoulder, all of v 1992 Oenopota (Buchozia) prisca (Desh.); King & which are characteristic of F. alienopriscus. Other Curry: 143, 145 (non Deshayes, 1862) (in part: Tilehurst approximately contemporaneous species of Pseudoliva Member, Crondall references only). resemble P. laudunensis in shape. Derivatio nominis: Named in honour of Prof. Dennis The German Priabonian species Fusulculus nodulosus Curry, who found all of the known examples of this species. (Beyrich, 1854) and F. pusillus (Beyrich, 1854) have also Holotype: BMNH PI TG 16682 (D. Curry coll.). been found in English Eocene sediments. Wrigley (1941: 164) united the two taxa, considering them to be simply Locus typicus: Harwich Formation, Tilehurst Member, variations within a single species, of which F. pusillus former brickyard south-west of Crondall, Hampshire. represented only a ribless form, although, pending Paratypes: BMNH PI TG 16683 – 16696 (D. Curry coll.), examination of specimens, we would regard F. nodulosus 14 examples from the type locality. and F. pusillus as specifically distinct, based on significant differences shown in the respective descriptions and figures Distribution: Early Eocene (Early Ypresian) of the of Beyrich (1854) and Koenen (1889). F. alienopriscus northern Hampshire Basin, England; known only from the differs from these two species in its more evenly fusiform, type locality and horizon. less sharply biconical shape, and by the presence of a Diagnosis : A small, rather narrowly fusiform Fusulculus

specimen height diameter aperture length aperture width Holotype PI TG 16682 3.2 2.0 1.9 0.9

Paratypes PI TG 16683 6.3 3.0 3.2 1.4

PI TG 16684 5.5 2.8 3.0 1.4

PI TG 16685 4.0 2.0 2.1 1.0

PI TG 16686 3.5 2.0 1.9 0.9

PI TG 16687 3.4 2.0 2.1 1.0

Table 2. Measurements (mm) of Fusulculus curryi sp. nov. 56 PACAUD & TRACEY EOCENE GASTROPOD 57 ornamented with slightly sinuous, thick axial ribs and spiral COSTA, E. M. DA 1776. Elements of Conchology or, an cords on base. Teleoconch whorls separated by shallow introduction to the knowledge of shells. London: 318pp., 7 pls. impressed sutures. Spiral sulcus indistinct. COX, L. R. 1960. Thoughts on the classification of the Gastropoda. Proceedings of the Malacological Society of London, Measurements (mm) seeTable 2. 33: 239-261. Description : Shell thick, solid, ovoid, consisting of c.5 CUVIER, G. 1797. Tableau élémentaire de l'histoire naturelle des whorls. Protoconch worn in all examples, c.1.5 whorls animaux. Paris: xvi + 710 pp., 14 pls. indicated. Teleoconch c. 3.5 convex whorls, with an DEFRANCE, F. 1826. Description (PORCE – PSY). In: indistinct thickened cord (bourrelet) below the suture. LEVRAULT (ed.) Dictionnaire des Sciences Naturelles. 53: 1- Ornament of low, sinuous axial ribs, extending abapically 544. Levrault, Strasbourg. of spiral sulcus which is shallow, probably terminating in a DESHAYES, G. P. 1835. Description des coquilles fossiles des triangular labial tooth (visible in the growth lines but worn environs de Paris. Levrault, Paris. 2(40-45): 495-780; Atlas 2: away from the lip of all known examples). Base below the pls.79-106. spiral sulcus ornamented with somewhat subdued spiral DESHAYES, G.P. 1862. Description des animaux sans vertèbres cords. Aperture ovate-elongate with broadly open siphonal découverts dans le Bassin de Paris. 2(29-32): 433-640; Atlas 2: notch. Columella simple, lip not thickened, regularly pls.27-39. Baillère, Paris. convex. D’ORBIGNY, A. 1850. Prodrome de Paléontologie Discussion: This new species is rather variable but the tall stratigraphique universelle des animaux mollusques et rayonnés, 2. narrow spire, and lack of both subsutural ramp and nodular Paris: 428 pp. shoulder, instantly separates it from F. alienopriscus sp. GREGORIO, A. DE 1880. Fauna di San Giovanni Ilarione nov. in particular, and from other described species in (Parisiano). Monografia, 1. Cefalopodi e Gasteropodi. Montaina, general. Although all the known examples are somewhat Palermo: 106pp., 9 pls. waterworn, their outline and proportions are more or less JEFFERY, P. & TRACEY, S. 1997. The Early Eocene London constant. Fusulculus priscus (Deshayes, 1862) has a more Clay Formation mollusc fauna of the former Bursledon sharply biconical shape, a short subsutural ramp on the last Brickworks, Lower Swanwick, Hampshire. Tertiary Research, 17: 75-137, pls. 1-15. whorl, a wider subsutural bourrelet, narrower, less pronounced axial ribs on the last whorl and spiral ornament KING, C. & CURRY, D. 1992. Molluscs from the Tilehurst over the whole whorl (Plate 1, fig.4). Member (London Clay Formation, Early Eocene) at Crondall and Up Nately (Hampshire). Tertiary Research, 13: 141-146. ACKNOWLEDGEMENTS KNIGHT, J. B., BATTEN, R. L., YOCHELSON, E. L. & COX, We would like to thank Daniel Ledon for the loan of the L. R. 1960. Supplement. Paleozoic and some Mesozoic juvenile examples of Pseudoliva laudunensis discussed Caenogastropoda and Opisthobranchia. In: MOORE, R. C. (ed.), herein, and David Ward for some of the digital Treatise on Invertebrate Paleontology, Part I, 1. Geological Society of America and University of Kansas Press, micrographs. The study was completed while one of us Lawrence, Kansas: I310-I324. (ST) was in receipt of a research grant from the University of Greenwich. KOENEN, A. von 1889. Das Norddeutsche Unter-Oligocän und seine mollusken-Fauna. 1: Strombidae, Muricidae, Buccinidae. Abhandlungen zur Geologischen Specialkarte von Preussen und REFERENCES den Thüringischen Staaten, 10: 1-280, pls. 1-23 BEYRICH, E. 1854. Die Conchylien des Norddeutschen LE RENARD, J. & PACAUD, J-M. 1995. Révision des Tertiärgebirges. Zeitschrift der Deutschen Geologischen Mollusques paléogènes du Bassin de Paris. 2: Listes des références Gesellschaft, Berlin, 6: 408-500, pls. 9-14. primaires des espèces - Cossmanniana, 3: 65-132. BROCCHI, G. 1814. Conchiologia fossile subappenina con PACAUD, J.-M. 1998. Nouvelles données sur le genre osservazioni geologiche sugli apennini e sul suolo adiacente. Popenoeum (Mollusca, Pseudolividae). Remarques taxinomiques Milano: 712pp., 16 pls. sur une espèce ubiquiste du Paléocène inférieur, Popenoeum BOUCHET, P. & VERMEIJ, G. J. 1998. Two new deep-water ambiguum (Binkhorst, 1861). Cossmanniana, 5: 1-30. 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Plate 1. Fig. 1 a, b. Fusulculus alienopriscus sp. nov., holotype BMNH PI TG 1111, Ypresian, London Clay Formation, division C1, Lower Swanwick, Hampshire; 3.6 x 2.3mm (from Jeffery & Tracey, 1997: pl.10, figs. 23, 24) Fig. 2. Fusulculus alienopriscus, sp. nov., juvenile with protoconch, BMNH GG 22551 (coll. S. Tracey), Ypresian, London Clay Formation, division C, Nursling, Hampshire; 1.38 x 0.93mm; arrow indicates lip of larval shell and start of first teleoconch whorl. Fig. 3 a, b. Fusulculus curryi sp. nov., holotype BMNH PI TG 16682 (coll. D. Curry), Ypresian, Harwich Formation, Tilehurst Member, Crondall, Hampshire; 3.2 x 2.0mm. Fig. 4 a, b. Fusulculus priscus (Deshayes) MNHN-LP R11783 (coll. J-M Pacaud) , Thanetian, Châlons-sur-Vesle (Marne), France; 5.2 x 2.6mm. Fig. 5 a, b. Pseudoliva laudunensis (Defrance), juvenile, MNHN-LP R11782 (coll. Ledon) Ypresian, Pourcy (Marne), France; 5.3 x 3.7mm. 58 PACAUD & TRACEY

Greenland and Denmark. Bulletin of the Geological Society of Denmark, 46: 53-67. RAVN, J. P. J. 1939. Etudes sur les Mollusques du Paléocène de Copenhague. Det Kongelige Danske Videnskarbernes Selskab, Biologiske Skrifter, 1: 1-103, pls.1-4. SALVINI-PLAWEN, L. V. & HASZPRUNAR, G. 1987. The Vetigastropoda and the systematics of streptoneurous Gastropoda (Mollusca). Journal of Zoology, London A 211: 747-770. SWAINSON, W. 1840. A Treatise on Malacology or the natural classification of shells and shell-fish. In: The Cabinet Cyclopaedia. London: 1-419. THIELE, J. 1929. Handbuch der systematischen Weichtierkunde, 1. Jena. 376pp. TRACEY, S. 1992. A review of the Early Eocene molluscs of Bognor Regis (Hampshire Basin), England. Tertiary Research, 13: 155-175. TRACEY, S., TODD, J. A., LE RENARD, J., KING, C. & GOODCHILD, M. 1996. Distribution of Mollusca in units S1 to S9 of the Selsey Formation (middle Lutetian), Selsey Peninsula, West Sussex. Tertiary Research, 16: 97-139, pls 1-3. TRAUB, F. 1979. Weitere Paleozän-Gastropoden aus dem Helvetikum des Haunsberges nördlich von Salzburg. Mitteilungen der Bayerischen Staatssamlung für Paläontologie und Historische Geologie, 19: 93-123, pls. 12-18. VERMEIJ, G. J. 1998. Generic revision of the neogastropod family Pseudolividae - The Nautilus, 111: 53-84, figs. 1-17.. VINCENT, G. 1878. Description de la faune de I'étage Landénien inférieur de Belgique. Annales de la Société Royale Malacologique de Belgique, 11: 111-160, pls. 6-10. WADE, B. 1917. New and little known Gastropoda from the Upper Cretaceous of Tennessee. Proceedings of the Academy of Natural Sciences of Philadelphia, 69: 280-304, pls.17-19. WRIGLEY, A. 1941. New species of Eocene Mollusca from the Isle of Wight. Proceedings of the Malacological Society of London, 24: 161-168, pl.11.

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