ABSTRACT. We have little understanding of the evolution of modern marine faunas from the European Atlantic coasts. In NW France very rich Neogene and early Quaternary mollusc assemblages occur that may shed light on the evolution of these modern faunas. Here we describe a Gelasian gastropod assemblage consisting of 76 species-level taxa from Selsoif (Manche, France), recovered from a sand extraction site between 1989-2020 by various collectors. The fauna reflects a range of intertidal to shallow subtidal palaeoenvironments. A few reworked Eocene fossils were also recovered. The extant portion of the Selsoif fauna is dominated by species with a Lusitanian signature, but also contains common species that nowadays have a Boreal-Celtic and Mediterranean signature. Furthermore, the fauna has quite some species in common with Early Pleistocene North Sea Basin faunas and also contains a few endemic species such as Calliostoma normanensis n. sp., Seila quinquecarinata n. sp., Alvania selsoifensis n. sp., Admete trigostomoides n. sp. The Selsoif fauna therefore represents a transitional fauna between Neogene and modern marine faunas of the European Atlantic.
KEYWORDS: Quaternary, Pleistocene, French Atlantic, gastropods, palaeoecology, biogeography, new species.
RÉSUMÉ. Nous comprenons peu l’évolution des faunes marines modernes des côtes atlantiques européennes. Dans le nord-ouest de la France, nous retrouvons des assemblages de mollusques très riches du Néogène et du début du Quaternaire, qui pourraient éclairer sur l’évolution de ces faunes modernes. Dans cet article, nous décrivons un assemblage de gastéropodes gélasiens composé de 76 taxons de Selsoif (Manche, France). La faune a été collectée par différentes personnes entre 1989 et 2020 sur un site d’extraction de sable. La faune représente des paléoenvironnements intertidal à subtidal peu profond, une très petite fraction de fossiles est remaniée de l’Éocène. La faune du Selsoif est dominée par des espèces à signature lusitanienne. Cependant, la faune contient des espèces ayant d’autres affinités, à savoir des espèces boréoceltiques à arctiques de l’Atlantique nord-est du Pléistocène inférieur, espèces européennes marines tempérées modernes et quelques espèces endémiques tel que Calliostoma normanensis n. sp., Seila quinquecarinata n. sp., Alvania selsoifensis n. sp., Admete trigostomoides n. sp. La faune Selsoif représente une faune de transition entre les faunes marines néogènes et modernes de l’Atlantique européen.
MOTS CLÉS: Quaternaire, Pléistocène, Atlantique francais, gastéropodes, paléocologie, biogéographie, espèce nouvelle.
Only very few areas in Atlantic Europe outside the southern North Sea Basin host Early Pleistocene marine fossil assemblages. Within the North Sea Basin, Middle Pleistocene mollusc faunas are almost entirely composed of extant species already showing a modern character ( Meijer et al. 2021 ; Wesselingh et al. 2023) . These ‘modern’ assemblages differ importantly from Pliocene-Early Pleistocene assemblages (e.g. Marquet 2001; Van Dingenen et al. 2015 ; Landau et al. 2020c ; Preece et al. 2020) . The lack of known Early Pleistocene faunas from the European Atlantic hinders our understanding of the evolution of the modern Atlantic marine biota. A well preserved Early Pleistocene fauna from Selsoif in Normandy (France) may enable us to fill this gap.
From the temperate Atlantic region very rich late Miocene and early Pliocene faunas are known, especially from the lower Loire region and adjacent Brittany (e.g., Brébion 1964; Lauriat-Rage 1982; Landau et al. 2020c) . However, these Neogene faunas bear very little resemblance to the modern coastal Atlantic faunas in terms of species and associations. Late Pliocene-Pleistocene faunas are known from the Channel region including faunas from the Wexford Gravels from southern Ireland (MacMillan 1964), and the St. Erth Beds of Cornwall (Harmer 1925; Mitchell et al. 1973) . However, these faunas have not been fully documented. The Pliocene-Early Pleistocene deposits from the Cotentin were described by Pareyn (1987) and Dugué et al. (2000 , 2007, 2009, 2012). Previously, they were classified as ‘upper Redonian’ (a regional French stage covering Tortonian to Gelasian marine intervals mostly centred around the lower Loire and Rennes region: Lauriat-Rage 1986; Van Dingenen et al. 2015) . However, Landau et al. (2020c) found that the Pleistocene faunas that they termed Redonian assemblage IV have very little in common with the Tortonian-Zanclean Redonian I-III assemblages.
In this study, we examined a particularly rich gastropod fauna recovered by different collectors during the past 40 years from a sand and gravel extraction site at Selsoif, located in Cotentin, France. Based on pollen analyses, the Selsoif fauna has been assigned an Early Pleistocene age (Pareyn 1987), and only a few species have been treated previously in taxonomic revisions (Marquet 2001; Van Dingenen et al. 2015 ; Hoeksema et al. 2020) . We present a complete taxonomic review of the gastropod assemblage in order to characterise the palaeoenvironmental settings of the Selsoif locality and improve our insights into the evolution of modern European temperate Atlantic coastal faunas.
The material used in this study was collected from a gravel and sand extraction site near Selsoif, municipality Saint-Sauveur-le-Vicomte, dept. Manche, France (Figs 1, 2). At the site, named Sablière du Grand Marais (49°21’20”N, 1°30’34”W), fossiliferous sands and gravels were dredged from 28-50 m water depth according to labels written by Arie W. Janssen. Shells were collected by handpicking and sieving of residues; details of sieving procedures are lacking. The collections were made by A.C. Janse (1995 and 1996), A.W. Janssen (1989) and W. Groeneveld (1995 and 1996). Complementary samples have been provided by S. Mermuys, collected at the same place in 2020 and an unsorted sample collected in 1995 by L. Vaessen.
We use the ecoregions defined by Spalding et al. (2007) but for brevity use the term “boreo-arctic ecoregion” for their Northern European Sea and Arctic ecoregions. Ecoregions used in this work from south to north are Mediterranean-Moroccan, French-Iberian (formerly Lusitanian), Boreal-celtic and Boreal-arctic (Fig. 3). For discussion on biogeographic terminology see Sacchetti et al. (2023 : 103).
The material used in this study is housed at Naturalis, collection Cainozoic Mollusca (indicated by RGM numbers), Collection of Stef Mermuys, Lansingerland (indicated by SM numbers) and in the Muséum national d’Histoire naturelle (MNHN), Paris, France (indicated by MNHN numbers). All species are briefly characterized, with longer discussions devoted to critical species and those that are new. We have restricted synonymy to major taxonomic sources and those most relevant to fossil assemblages. The nomenclature follows WoRMS, 2023 with deviations discussed below.
The Cotentin region is located at the transition of the Palaeozoic Armorican Massif in the west and the Paris Basin in the east ( Dugué et al. 2000 : fig. 1). The region contains a number of small sediment-filled grabens ( Baize et al. 1998) resulting from Alpine orogenesis creating fault and deformation in the Channel region. The basin fill occurred during three transgressive intervals (middle Eocene, Miocene and Plio-Pleistocene) and is dominated by marine deposits. The three phases are separated by periods of non-deposition and erosion signifying emergent phases (Paleocene, middle Oligocene to Early Miocene and Middle Miocene to Early Pliocene: Dugué et al. 2007) . Most successions are found in the subsurface, outcrops are rare. The Cenozoic stratigraphic sequence was studied after a drilling campaign in the 1980s and discussed by various authors (e.g., Pareyn 1987; Dugué et al. 2007 ; 2012; Baize et al. 1998 ). Subdivision and correlations with other NW European formations were proposed based on different faunal groups (Lauriat-Rage 1982 for bivalves; Kasimi 1986 for ostracods and Le Calvez 1987 for foraminiferans). Further age constraints were provided by palynology (Pareyn 1987). The Cotentin Basin is separated into two large subbasins, the east-west oriented Saint-Sauveur-le-Vicomte subbasin in the north, and the north-south oriented Carentan subbasin in the south (Pareyn 1987). Selsoif is located in the centre of the Saint-Sauveur-le-Vicomte subbasin. The entire Pliocene-Pleistocene sequence contains six formations that represent a transition of marine to fluvial dominated settings ( Dugué et al. 2012) . Exact age estimates for the formations are scarce and uncertain. The oldest formation is the Late Pliocene Grès Coquillier de Marchésieux (thickness of around 70 m), which represents deep marine settings. It is known only from the subsurface around Marchésieux ( Dugué et al. 2000) in the southern subbasin. The overlying Marne du Bosq d'Aubigny (thickness of around 55 m) is a fossil-rich marly unit whose age is estimated between 2.75 and 2.4 Ma ( Van Dingenen et al. 2015 ; Landau et al. 2020c) . Intercalated shell intervals, the so-called Falun de Bohon (maximum thickness of 50 m), are present around Saint Georges de Bohon forming the top of the Marne du Bosq d'Aubigny. In the region of Saint-Sauveur-le-Vicomte, the marly and shelly intervals appear intermixed ( Baize et al. 1998) . The overlying Marnes the Saint Nicolas de Pierrepont Formation (maximum thickness 40 m) is only known from boreholes ( Dugué et al. 2000) . That unit has a large extension ranging from Saint-Sauveur le Vicomte in the north to the south part of the Cotentin Basin ( Baize et al. 1998 ). Overlying is the Sables de Saint Vigor Formation (maximum thickness 30 m) that is present throughout the Cotentin Basin and lacks fossils, making it difficult to determine its age. It has been referred to as the lowermost Pleistocene unit ( Dugué et al. 2000) . Locally, the upper part of the Sables de Saint-Vigor is eroded, and channels are filled with so-called puddingstone and covered by fluviatile deposits of the Sables de la Lande de Millière Formation ( Dugué et al. 2000) . After the early Pleistocene, the region became uplifted and (marine) deposition ceased.
The sequence contains two transgressive-regressive cycles covering the Pliocene-Pleistocene transition. The oldest maximum transgression occurred during deposition of the Grès Coquillier de Marchésieux during the Late Pliocene. The second transgressive maximum was reached at the transition of the Marnes de Saint-Nicolas-de-Pierrepont Formation and the Sables de Saint Vigor Formation (Dugué et al. 2009) . The fossiliferous intervals of Selsoif are part of the Saint-Sauveur-de-Pierrepont Formation of Pareyn (1987) that was subsequently renamed as Saint-Nicolas de Pierrepont Formation (Dugué et al. 2000) . The stratigraphic age of the Selsoif succession is not well constrained. Pareyn (1987) correlated it with Tiglian-C based on palynological analyses of Clet-Pellerin (1983). The Tiglian is a regional Gelasian stage defined for the southern North Sea Basin mostly based on pollen analysis. However, the original Tiglian pollen zones likely represent a number of recurring glacial-interglacial successions whose attribution to marine isotope stages (MIS stages) is not resolved (Westerhoff et al. 2020 ; see Discussion). However, the Selsoif association is of a Gelasian age.
WG W. Groeneveld;
ACJ A. C. Janse;
AWJ A. W. Janssen;
LV L. M. B. Vaessen;
SM S. Mermuys;
The gastropod fauna of Selsoif is composed of 76 gastropod taxa (Table 1) representing mostly a shallow marine warm temperate fauna.
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(Fig. 4 A1, A2)
Maximum height 9.1 mm, diameter 28.2 mm. — RGM.1364897 (2), leg. WG; RGM.1364961 (5), leg. ACJ; RGM.1365058 (18), leg. AWJ; RGM.1365190 (1), leg. AWJ.
Low patelliform shell; apex placed two-thirds distance from anterior margin. Sculpture of about 34 low, narrow, scabrous primary ribs with single secondary intercalated in interspaces. Edge finely crenulated.
Fossils records are scarce. Pliocene (indeterminate): Calabria and Modena, Italy ( Bucquoy et al. 1886 : 471). — Early Pleistocene: Atlantic, Selsoif, France (this paper). — Middle/Late Pleistocene: Italy ( Buccheri et al. 2014) . Today this species occurs in the Atlantic south of Bay of Biscay, Strait of Gibraltar, to the coast of North Africa, including the Macaronesian Islands ( Zegaoula et al. 2016) , and into the Mediterranean Sea from Spain ( Gofas et al. 2011) to the Aegean Sea ( Öztürk et al. 2014) . It inhabits upper midlittoral rocky environments ( Bensettiti et al. 2004 : 112).
Separation of et al. 1996) . The specimens from Selsoif probably represent et al. 1996) .
(Fig. 4 B1, B2)
Maximum diameter 15.1 mm. — RGM.1310830 (1), leg AWJ; RGM.1315059 (1), leg AWJ.
Intermediate-high patelliform shell; apex placed two-thirds distance from anterior margin. Largest width at one-thirds distance from anterior margin. Sculpture of about 28 robust primary ribs that can develop slightly irregular knobs, with low secondary ribs and very fine but irregularly developed and spaced tertiary ribs. Edge coarsely wavy, corresponding mostly to primary ribs.
Lower Pleistocene: Atlantic, Selsoif, France (this paper). — Upper Pleistocene: Channel region, Portland and Selsey (Harmer 1921). — Upper Pleistocene-Holocene: Netherlands (Wesselingh & Pouwer 2011). — Present-day: NE Atlantic coast from Norway to Gibraltar. Rarely reported from western Mediterranean.
As discussed above, identification of fresh
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(Fig. 4 C1, C2)
et al. 2003 : 8, pl. 5, fig. 3.
et al. 2017 : 79, pl. 2, figs 1-2.
For more, see synonymy list in Landau et al. (2003 ; 2017)
Maximum height 8.4 mm, width 2.9 mm. — RGM.1364898 (15), leg. WG; RGM.1364962 (236), leg. ACJ; RGM.1365189 (1), leg. ACJ; RGM.1365060 (54), leg. AWJ.
Shell patelliform, elongate-ovate in profile, apex placed just posterior to centre to close to posterior margin; surface smooth, ribbing suggested in figured specimen due to erosion of surface.
Middle Miocene: Atlantic of Loire Basin, France (Glibert 1949). — Upper Miocene: Atlantic, NW France ( Landau et al. 2017) . — Lower Pliocene: Coralline Crag, England (Harmer 1925); central Mediterranean, Italy, (Chirli 2004), Tunisia (Fekih 1975). — Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1925); Atlantic, Mondego Basin, Portugal (Silva 2001); western Mediterranean, Estepona Basin, S. Spain ( Landau et al. 2003) ; central Mediterranean, Italy (Sacco 1896a). — Lower Pleistocene: Atlantic, Selsoif, France (this paper); central Mediterranean, Italy (Cerulli-Irelli 1916; Ruggieri & Greco 1965; Taviani et al. 1998) ; eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Upper Pleistocene: NSB, Netherlands (Wesselingh & Pouwer 2011); Atlantic, England (Harmer 1925); western Mediterranean, Balearic Islands (Cuerda Barceló 1987). — Holocene: NSB (Strand Petersen 2004). Today this species occurs in the Atlantic, from northern Scandinavia to Cape Verde Islands at 0-100 m depth attached to hard substrates (Poppe & Goto 1991) and in the Mediterranean ( Giannuzzi-Savelli et al. 1994) from the Spain ( Gofas et al. 2011) coast to the Aegean Sea ( Öztürk et al. 2014) .
This species is extremely variable in height and profile and displays few distinct shell characters. Colour pattern is not preserved in the Selsoif material. For further discussion see Landau et al. (2003 , 2017).
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(Fig. 4 D1, D2)
et al. 2003 : 26, pl. 4, fig. 4; 2017: 80, pl. 3, figs 1-4. — Chirli & Linse 2011: 35, pl. 4, fig. 5. — Ceulemans et al. 2016 : 54, pl. 1, figs 5, 6. — Chirli & Forli 2017: 24, fig. 5a.
For more, see synonymy list in Landau et al. (2003 , 2017) and Ceulemans et al. (2016) .
Maximum height 23.5 mm, diameter 11.9 mm. — RGM.1364899 (3), leg. WG; RGM.1364964 (7), leg. ACJ; RGM.1365061 (24), leg. AWJ; RGM.1365174 (1), leg. AWJ; RGM.1365062 (1), leg. AWJ.
Patelliform shell with ovate profile; apical perforation two-third distance from anterior margin; sculpture of primary to tertiary ribs differentiated to a variable degree.
Middle Miocene: Paratethys (Strausz 1966; Bałuk 1975). — Upper Miocene: NW France ( Ceulemans et al. 2016) ; Proto-Mediterranean (Sacco 1896a). — Lower Pliocene: NW France ( Ceulemans et al. 2016) , Coralline Crag, England (Harmer 1923), Belgium (Marquet & Landau 2006); central Mediterranean (Chirli 2004). — Upper Pliocene: Red, Crag, England (Harmer 1923); western and central Mediterranean ( Landau et al. 2003 ; Malatesta 1974; Caprotti 1976; Cavallo & Repetto 1992); France (Chirli & Richard 2008). — Lower Pleistocene: NW France (Brébion 1964), Selsoif (this paper); central Mediterranean (Cerulli-Irelli 1916; Malatesta 1960; Chirli & Forli 2017); eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Upper Pleistocene: England (Harmer 1923), Netherlands (Wesselingh & Pouwer 2011); western Mediterranean, Balearic Islands (Cuerda Barceló 1987). Today this species occurs from the Atlantic British Isles to Canaries, Mediterranean and Black Sea on rocky shores under stones and rocks, especially where a little silt occurs, living near sponges, on which it feeds (Poppe & Goto 1991).
This species is highly variable in height and the differentiation between the primary to tertiary ribs (see discussion in Ceulemans et al. 2016 : 55). Some of the Selsoif specimens have weaker primaries than seen in some Mediterranean fossil material.
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(Fig. 4 E1, E2)
et al. 2016 : 57, pl. 2, fig. 2. — Brunetti & Cresti 2018: 30, fig. 19.
For more, see synonymy list in Ceulemans et al. (2016) .
Maximum height 8.3 mm, diameter 11.7 mm. — RGM.1364900 (2), leg. WG; RGM.1364901 (5), leg. WG; RGM.1365175 (1), leg. WG, RGM.1364965 (13), leg. ACJ; RGM.1365376 (1), leg. ACJ; RGM.1364966 (73), leg. ACJ; RGM.1364967 (28), leg. ACJ; RGM.1365063 (13), leg. AWJ; RGM.1365064 (13), leg. AWJ; RGM.1365065 (22), leg. AWJ; RGM.1365176 (1), leg. AWJ.
Elevated conical shell; apex placed about two thirds distance from anterior margin, sculpture of alternating primary and secondary ribs, slightly weaker concentric cords forming fine reticulated surface pattern; selenizone narrow, slit extending to one-third shell height.
Middle Miocene: Loire Basin, France (Glibert 1949). — Upper Miocene: NW France (Brébion 1964; not confirmed by Landau et al. 2017) . — Lower and Upper Pliocene: NSB, England (Wood 1848; Harmer 1923), Belgium (Marquet 1998), The Netherlands (Beets 1946); Atlantic, NW France ( Ceulemans et al. 2016) , Portugal (Silva 1990, 2001); western and central Mediterranean (Fekih 1975; Piani 1984; Cavallo & Repetto 1992; Landau et al. 2003 , 2011; Brunetti & Cresti 2018). — Lower Pleistocene: Atlantic, NW France (Brébion 1964), Selsoif, France (this paper); eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Pleistocene (indeterminate): central Mediterranean, Italy (Sacco 1896a; Cerulli-Irelli 1916). Today it occurs along the European Atlantic coasts from Scandinavia to the Mediterranean, at a depth between 0 and 700 m (Piani 1984).
For discussion on the shell variability and comparison with congeners see Ceulemans et al. (2016 : 57).
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(Fig. 4 F1-F3)
et al. 2003 : 42, pl. 9, fig. 6. — Chirli 2004: 78, pl. 32, figs 10-12, pl. 33, figs 1-4. — Chirli & Linse 2011: 39, pl. 6, fig. 1. — Chirli & Forli 2017: fig. 6b.
et al. 2017 97, pl. 21, figs 1-4.
For more, see synonymy list in Landau et al. (2003) .
Maximum height 9.4 mm, width 7.3 mm. RGM.1364903 (36), leg. WG; RGM.1364969 (48), leg. ACJ; RGM.1365178 (1), leg. ACJ; RGM.1365067 (60+) leg. AWJ.
Shell trochiform; protoconch severely worn or not preserved. Teleoconch of six flat-sided whorls separated by superficial suture; sculpture of five spiral cords, abapical one broader, stronger, placed just above the suture; fine lamellar ribs visible in cord interspaces giving them a pitted appearance; base flattened bearing six concentric cords; aperture subquadrate; columella short, slightly oblique bearing subobsolete fold.
?Upper Miocene: NW France (Brébion 1964; conspecificity uncertain; Landau et al. 2017) . — Pliocene: Italy (Brunetti & Vecchi 2005). — Upper Pliocene. Western Mediterranean, S. Spain ( Landau et al. 2003) . — Lower Pleistocene: Atlantic, southern England (Harmer 1923), Selsoif, France (this paper); central Mediterranean, Italy (Chirli & Forli 2017); eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Upper Pleistocene: Atlantic, Santa Maria Island, Azores (Callapez & Ferreira 2000); western Mediterranean, Balearic Islands (Cuerda Barceló 1987). Today, this species occurs in the Atlantic from the west coast of Scotland to the Canary Islands and Azores (Fretter & Graham 1977; Hernández et al. 2011) , Mediterranean Sea (Manousis 2021) and Aegean Sea ( Öztürk et al. 2014 ).
(Fig. 4 G1-G3)
et al. 2003 : 44, pl. 10, fig. 1. — Chirli 2004: 81, pl. 33, figs 9-16. — Pouwer & Wesselingh 2012: 159, figs 27-29.
Maximum height 7.0 mm (incomplete), width 7.2 mm. — RGM.1365068 (1), leg. AWJ; RGM.1365179 (1), leg. AWJ.
Shell trochiform; protoconch and early teleoconch whorls not preserved; last two whorls weakly convex, separated by moderately deeply impressed suture; sculpture of seven to eight irregular, flattened spiral cords; axial sculpture of fine, close-set axial growth lamellae present only in the interspaces; base flattened bearing seven concentric cords; columella short, oblique, curved, bearing stout columellar fold delimiting siphonal fasciole.
Lower Pliocene: central Mediterranean, Italy (Chirli 2004). — Upper Pliocene: western Mediterranean, Estepona Basin, Spain ( Landau et al. 2003) ; central Mediterranean, Italy (Caprotti 1976; Cavallo & Repetto 1992). — Upper Pleistocene: southern North Sea Basin (Pouwer & Wesselingh 2012). — Present-day: Atlantic, Orkneys south into the western Mediterranean, 5-200 m deep, mainly on gravel bottoms (Poppe & Goto 1991).
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(Fig. 4 H1-H3, I1-I3)
? non
partim, not pl. 58, figs 25, 26) (Upper Pleistocene records only).
et al. 2020 : 70, pl. 50.
et al. 2017 : 804, fig. 5.
Maximum height 12.7 mm, width 12.4 mm. — RGM.1364902 (64), leg. WG; RGM.1364968 (125), leg. ACJ; RGM.1365177 (1), leg. ACJ; RGM.1310831 (1), leg. ACJ; RGM.1365066 (146), leg. AWJ.
Shell low trochiform; five weakly shouldered teleoconch whorls separated by narrowly impressed suture; sculpture of numerous, fine, irregular spiral cords; axial sculpture reduced to prosocline growth-lines; base depressed bearing numerous concentric cords; umbilicus well developed, moderately narrow and deep; columella short, straight, oblique; colour pattern of red-orange zigzag flammules preserved in all specimens.
(Late Pliocene-Early Pleistocene: NSB and Irish Sea [ Wood 1848; Harmer 1923 ] likely based on misidentifications [ Pouwer & Wesselingh 2012 ]). Lower Pleistocene: Atlantic, Selsoif, France (this paper). Upper Pleistocene: British Isles (Harmer 1923). — Holocene: NSB (Strand Petersen 2004). Today this species is predominantly found in cooler waters in the NSB and along the European Atlantic coasts North Norway, Iceland to Gibraltar (Fretter & Graham 1977) and adjacent western Mediterranean ( Giannuzzi-Savelli et al. 1994) . Records south of Gibraltar in the Atlantic and into the Mediterranean, such as south Manousis (2021), need to be confirmed.
Based on molecular data, this species is now placed in the genus et al. 2017) . Most specimens from Selsoif are small and depressed (Fig. 4 I1-I3). One large, gerontic specimen (Fig. 4 H1-H3) is taller spired, with more strongly shouldered whorls, the shoulder marked by two slightly strengthened cords. In general, the Selsoif specimens have slightly more shouldered whorls than extant specimens, but we consider them conspecific.
(Fig. 4 J1-J3)
partim, pl. 59, fig. 11, not pl. 59, fig. 10) (Upper Pleistocene-Recent records only).
noninEiríksson & Símonarson, 2021).
Height 4.8 mm. — RGM.1310832 (1), leg. SM; n.n. (8), leg. SM.
This small
Lower Pleistocene: Atlantic, Selsoif, France (this paper). — Late Pleistocene: Channel region, Selsey, west coast of Scotland, Largo bay, Belfast, Livorno (Italy) and Christiana Fjord and Trondhjem in Norway (Harmer 1923). — Recent: Iceland and northern Norway to Gibraltar (Pouwer & Wesselingh 2012; Harmer 1923).
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urn:lsid:zoobank.org:act:662E6CB4-0935-4328-8183-A0419A765952
MNHN.F.A91315, height 12.8 mm (incomplete), width 14.7 mm, leg. AWJ.
Trochoid shell with broad, evenly conical spire, flattened base, imperforate, spiral sculpture of four finely beaded primary cords with secondary threads intercalated, 13 weakly beaded cords on base, columella strongly oblique.
Named after the Region of Normandy in which the type locality is situated.
Selsoif, Saint-Sauveur-le-Vicomte, Manche department, NW France.
Gelasian, Early Pleistocene.
Maximum height approx. 17 mm (incomplete), width 16.4 mm. — RGM.1364904 (11 juveniles), leg. WG; RGM.1364970, (27 juveniles + one incomplete adult), leg. ACJ; RGM.1365069 (22 incomplete adults and fragments), leg. AWJ.
Lower Pleistocene: Atlantic, Selsoif, France (this paper).
Shell medium sized, trochiform; protoconch not preserved; five teleoconch whorls preserved, initially flat-sided, later whorls slightly swollen just above suture, separated by superficial suture; sculpture on first teleoconch whorl of four finely beaded spiral cords; on second whorl, single secondary spiral intercalated between adapical spiral and second primary cord, strengthening abapically to become almost equal in strength to primaries on later adult whorls. On penultimate whorl secondaries appear between the second and third primaries and between the fourth and the suture. Axial sculpture reduce to weak prosocline growth-lines. Last whorl 47% of total height; slightly inflated and strongly rounded at periphery; on last whorl single secondary intercalated in all interspaces, some secondaries almost equal in strength to primaries. Base depressed, imperforate, bearing 13 weakly beaded concentric cords, widening and strengthening toward columella. Aperture subquadrate, 35% of total shell height; outer lip simple, strongly prosocline in profile, roundly angled at base. Columella strongly inclined abaxially, slightly thickened, parietal callus absent.
The Selsoif material has sculpture similar to that of et al. (1989 : pl. 8, figs 5, 6) from the Lower Pliocene of La Limouzinière (NW France). Several similar
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(Fig. 5 B1-B3)
et al. 2016 : 73, pl. 10, fig. 1. — Landau et al. 2017 : 142, pl. 66, fig. 1.
For more, see synonymy list in Ceulemans et al. (2016) .
Maximum height 5.4 mm, width 3.9 mm. — RGM.1365319 (47), leg. AWJ; RGM.1365303 (1), leg. AWJ; RGM.1365300 (2), leg. AWJ; RGM.1365301 (29), leg. AWJ; RGM.1365302 (15), leg. AWJ.
Small depressed skeneimorph shell; protoconch of one smooth whorl; teleoconch of two whorls separated by deep suture; sculpture of fine spiral cords that continues inside the deep umbilicus.
Upper Miocene: Atlantic, NW France ( Landau et al. 2017) . — Lower Pliocene: NSB, Luchtbal Member, Belgium (Marquet & Landau 2006); Atlantic, NW France ( Ceulemans et al. 2016) . — Lower Pleistocene: Atlantic, Selsoif, France (this paper). Today this species is present from the Atlantic SW England (Fretter & Graham 1977) to southwards into the Mediterranean, Corsica ( Van Aartsen et al. 1984) .
There is some variation in shape in the specimen from Selsoif, some slightly more depressed than others and in some specimens the last portion of the last whorl is slightly disjunct. fide Rubio et al. 2004 : 131) but the protoconch is rugose rather than smooth.
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(Fig. 5 C1, C2)
Maximum height 0.61 mm, width 0.96 mm.
RGM.1362824, height 0.6 mm, width 0.96 mm, leg. AWJ.
Paratype 1, RGM.1362825, height 0.61 mm, width 0.95 mm, leg. AWJ; paratype 2, RGM.1362630 (8), leg. AWJ.
“Shell very small, wider than high, rather solid, opaque, depressed skeneiform, up to 2.25 whorls protoconch included. Suture deep, canaliculate. Protoconch paucispiral, ¾ whorls, smooth, D about 0.30 mm. Protoconch-teleoconch border delimited by shallow groove. Aperture prosocline and circular. Peristome sharp, simple, almost flat, continuous. Umbilicus open, deep, bordered by sharp peri-umbilical edge (Figs 1c, 1e, 2c, 2e). Teleoconch sculptured with ripple-like, prosocline, straight to sigmoid, flat-topped, partly dichotomous axial ribs, 40-50 on last whorl, broader than interspaces; the branched rib-parts at the periphery coalesce adapically at a short distance from the penultimate whorl and abapically on the base. Body whorl depressed, convex, weakly angled at shoulder ( Hoeksema et al. 2020 : figs 1d, 2d) and in two spirals around the peri-umbilical edge ( Hoeksema et al. 2020 : figs 1e, 2e). Dimensions: H up to 0.65 mm, W up to 1.00 mm.” ( Hoeksema et al. 2020 : 6).
This species is known only from Selsoif. For full discussion see Hoeksema et al. (2020 : 6).
Lower Pleistocene: Atlantic, endemic to Selsoif ( Hoeksema et al. 2020) .
-
-
(Fig. 5 D1-D3)
et al. 2017 : 153, pl. 77, figs 1-6.
For more, see synonymy list in Landau et al. (2017) .
Height 2.3 mm (incomplete), width 2.0 mm. — RGM.1365082 (1), leg. AWJ.
Shell turbiniform, only last whorl showing a flattened profile to the upper half, rounded periphery, bearing reddish zigzag colour pattern interrupted by two rows of white blotches the upper placed below the suture, the lower mid-whorl.
Upper Pliocene: Atlantic, St Erth, England (Harmer 1925), Selsoif, NW France (this paper). Today it is present from the western coasts of British Isles (absent in Scandinavia and North Sea) south to Gibraltar (Fretter & Graham 1977). Further south it is replaced by the subspecies et al. 2011) . It lives in rocky pools, 0-35 m depth, common on red weeds (Fretter & Graham 1977).
Two subspecies occur today. et al. (2017 : 153) noted that all forms occurred together in those assemblages and postulated that these subspecies may not have separated at the time. The material from Selsoif is scant, but suggest the Atlantic form had split by at least the Gelasian.
-
-
(Fig. 5 E)
et al. 2016 : 65, pl. 5, fig. 15. — Landau et al. 2018 : 246, pl. 74, fig. 1.
Maximum height 4.0 mm, width 8.2 mm. — RGM.1364906 (10), leg. WG; RGM.1364971 (17), leg. ACJ; RGM.1365071 (100+), leg. AWJ; RGM.1365216 (1), leg. AWJ.
Irregularly coiled gastropod with round cross-section; surface details worn in all material and protoconch lacking.
Lower Miocene: Proto-Mediterranean, Italy (Sacco 1896b). — Lower-Middle Miocene: NSB, Belgium (Glibert 1952), Germany (Anderson 1964; Moths 1989), Netherlands (Janssen 1984). Middle Miocene: Atlantic, Aquitaine Basin, (Cossmann & Peyrot 1924; Lozouet et al. 2001) , Loire Basin, France (Glibert 1949); Paratethys, Poland (Friedberg 1914; Bałuk 1970, 1975), Austria (Hörnes 1856; Tejkal et al. 1967 ; Schultz 1998), Bulgaria (Kojumdgieva & Strachimirov 1960), Hungary (Strausz 1966), Romania (Moisescu 1955; Stancu & Andreescu 1968); Proto-Mediterranean, Karaman Basin, Turkey (Erünal-Erentoz 1958; Landau et al. 2013) . — Upper Miocene: Atlantic, NW France ( Landau et al. 2018) ; Proto-Mediterranean, Italy (Sacco 1896b), Tunisia (Stchepinsky 1938). — Lower Pliocene: NSB, England (Wood 1848; Harmer 1918), Belgium (Glibert 1958; Marquet 1998); Atlantic, NW France ( Van Dingenen et al. 2016) , Spain (González-Delgado 1986; Landau et al. 2011) , Morocco (Lecointre 1952); western Mediterranean, NE Spain, (Martinell & Domènech 1984; Solsona 1998), France (Fontannes 1879); central Mediterranean, Italy (Sacco 1896b; Palla 1967; Caprotti 1974; Anfossi et al. 1983 ; Baroncelli 2001); Tunisia (Fekih 1975). — Lower-Upper Pliocene: Atlantic, Mondego Basin, Portugal (Zbyszewski 1959; Silva 2001); western Mediterranean, Spain ( Landau et al. 2004) ; central Mediterranean, Italy (Malatesta 1974; Cavallo & Repetto 1992). — Upper Pliocene-Pleistocene: NW France (Brébion 1964). — Lower Pleistocene: Atlantic, Selsoif (this paper); central Mediterranean, Italy (Cerulli-Irelli 1912; Taviani et al. 1998) .
Specific identity is difficult to establish due to lack of protoconch in the studied material. Here tentatively assigned to , a common species in fossil faunas from North West Europe. The modern species et al. (2018 : 246).
-
-
(Fig. 5 F1-F3)
Maximum height 6.1 mm, width 1.8 mm. — RGM.1364907 (2), leg. WG; RGM.1364972 (46), leg. ACJ; RGM.1365268 (1), leg. ACJ; RGM.1365072 (70), leg. AWJ; RGM.1365284 (1), leg. AWJ; RGM.1365375 (1), leg. AWJ.
Robust turriform shell with pronounced suture. Broad paucispiral protoconch of two smooth whorls. Teleoconch whorls with three spirals cords and slightly prosocline axials ribs forming low tubercles at intersections.
Lower Pliocene: NSB, Kattendijk Formation, Belgium (Glibert 1958), Oosterhout Formation, the Netherlands (RGM collections). — Lower Pleistocene: Atlantic, Selsoif, France (this paper).
The Selsoif specimens have more robust axials and lower tubercles that make it slightly different from the specimens illustrated by Glibert (1958) that tend to have finer ribs and less pronounced knobs. Otherwise, the number and organisation of the ribs and cords of the Early Pliocene North Sea Basin (NSB) specimens and the Selsoif material are identical. The geographical and temporal distribution gap between the NSB Early Pliocene and French Gelasian occurrences is remarkable. A et al. (2012 : 42) is not substantiated.
(Fig. 5 G1-G3)
et al. 2004 : 10, pl. 2, fig. 1. — Wesselingh et al. 2012 : 39, figs 6-9. — Chirli & Forli 2017: 43, fig. 12f.
For more, see synonymy list in Landau et al. (2004) .
Maximum height 3.8 mm, width 1.3 mm. — RGM.1365269 (1), leg. AWJ; RGM.1365073 (3), leg. AWJ.
Robust
Upper Pliocene: western Mediterranean, S. Spain ( Landau et al. 2004) . — Lower Pleistocene: Selsoif, NW France (this paper); central Mediterranean, Italy (Chirli & Forli 2017). — Middle Pleistocene: NSB, the Netherlands ( Wesselingh et al. 2023) . — Upper Pleistocene: NSB, the Netherlands (RGM collections); western Mediterranean, Balearic Islands (Cuerda Barceló 1987). Today it occurs from the North Sea to North Africa, and into the Mediterranean (Poppe & Goto 1991)
Currently WoRMS (2023) considers et al. (2004) , the main differences between the two species are that et al. 2012 ; 2023). The species et al., 2018
-
-
(Fig. 5 H1-H3, I1-I3)
et al. 2016 : 117, pl. 3, fig. 1.
et al. 2018 : 27, figs 6-8.
For more, see synonymy list in Van Dingenen et al. (2016) .
Maximum height 42.4 mm, width 11.5 mm. — RGM.1365074 (700+) leg. AWJ; RGM.1365285 (1), leg AWJ, RGM.1310833 (1), leg. AWJ; RGM.1364908 (140+) leg. WG; RGM.1364974 (200+) leg. ACJ; RGM.1365085 (4), leg WG.
Medium sized
Lower Pliocene: NSB, Coralline Crag Formation, England (Wood 1848; Harmer 1918); Atlantic, NW France ( Van Dingenen et al. 2016) . — Upper Pliocene: NSB, Red Crag Formation, England (Wood 1848; Harmer 1918); Oorderen and Kruisschans Members, Belgium (Marquet 1997, 1998), Oosterhout Formation, The Netherlands ( Moerdijk et al. 2018) . — Upper Pliocene: Atlantic, St Erth, England (Harmer 1918), Selsoif, NW France (this paper).
Marwick (1957) highlighted the importance in the order of appearance of the cords on the neanic whorls and used a lettering system for the spiral cords in which B was the medial primary, D the peribasal primary generally involved with the suture, A was the first to appear adapical to B and C the first to appear abapical to B, between B and d (Marwick 1957: 148), and the shape of the lateral and basal sinuses. Subsequently, this work has been neglected by almost all European workers. More recently, Harzhauser & Landau (2019) applied Marwick’s model and proposed generic attributions for European fossil turritellids from the Paratethys. Following the diagnostic generic characters proposed in that work, the Selsoif specimens are readily attributed to
Based on the whorl profile and adult sculpture we attribute these specimens to et al. 2016) or from the Pliocene Coralline and Red Crag Formations of eastern England, and Pliocene units from Belgium and the Netherlands (Harmer 1925; Marquet 1998; Moerdijk et al. 2018) , the Selsoif specimens tend to be straight-ided to slightly concave with fine slightly irregular spiral cordlets. The relative smoothness of the later whorls is reminiscent of the endemic Early Pliocene NSB e.g., Van Regteren Altena et al. 1955 ) are not supported by in situ findings (FW pers. observation) and are not included in the distribution.
-
-
(Fig. 5 J1-J3)
cf.
cf.
cf. et al. 2016 : 162, pl. 14 figs 7, 8.
Maximum height 5.8 mm (incomplete), width 3.2 mm. — RGM.1364975 (3), leg. ACJ; RGM.1365280 (1), leg. AWJ.
Apex lacking on all material. Overall shell is slightly spindle shaped. Adult whorls separated by superficial suture; sculpture of three spiral ribs, middle rib slightly stronger than other ribs; numerous weak, prosocline ribs, with rounded tubercles developed at intersections; on last whorl the fourth basal rib is smooth to slightly knobbed and a fifth rib in one specimen is smooth reminiscent of a recent Canarian Island morph of this species (H. Bakker, pers. comm.). The Selsoif specimens are large for this species.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). Today
The three extant West European species
-
(Fig. 6 A1-A3)
Height 3.7 mm (incomplete) mm, width 1.5 mm. — RGM.1365279 (1), leg. AWJ.
We only have one damaged specimen lacking protoconch. The earliest whorl preserved has two primary spiral cords, S1 and S3 with large tubercles developed at intersections with narrow axial ribs; the profile between the two spiral cords is concave. S2 appears between S1 and S2, strengthening on later whorls, of almost equal strength on last whorl. The overall shape and tardive S2 fits well in the genus
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
-
-
(Fig. 6 B1-B4)
Maximum height 4.2 mm, width 1.5 mm .Paratypes RGM.394128 (2), leg. AWJ. Other material: RGM.1364909 (1), leg. WG; RGM.1364978 (90+) leg. ACJ; RGM.1365180 (1), leg. ACJ; RGM.1365075 (100+) leg. AWJ.
“Shell small, turriculate, rather broad, with an apical angle of more than 25°, consisting of eight whorls. Protoconch paucispiral, large (broader than first teleoconch whorl), smooth, very tumid, consisting of only one whorl. Teleoconch whorls tumid for genus. Sculpture starting with axial ribs and two abapical spirals. Third, adapical spiral starting one whorl later and always remaining weaker than both other spirals. Spiral ribs about as broad as intercostal areas. Axial ribs strong, clearly delimited, about as broad as the intercostal spaces, widely spaced, 24 to 30 in number on the middle whorls. Suture deep. Shell base smooth, convex, without additional spirals; aperture nearly square; siphonal canal rather long for genus”. (Marquet 2001: 196).
Lower Pleistocene: Atlantic, endemic to Selsoif, NW France (Marquet 2001).
This endemic species is quite unlike its European Atlantic and NSB congeners in being much squatter and having a wider apical angle. For full discussion see Marquet (2001: 196).
(Fig. 6 C1-C4, D1-D3)
urn:lsid:zoobank.org:act:728DCCE3-1D4E-4A21-B87B-CD41D41CAA98
MNHN.F.A91316, height 8.1 mm (incomplete), width 2.7 mm, leg. AWJ.
Maximum height 8.1 mm (incomplete), width 2.7 mm. — RGM.1365119 (26 fragments), leg. AWJ; RGM.1365015 (14 fragments), leg. ACJ; RGM.1364931 (1), leg. WG.
Named reflecting the spiral sculpture consisting of five cords on later adult whorls.
Selsoif, Saint-Sauveur-le-Vicomte, Manche department, NW France.
Gelasian, Early Pleistocene
Small
Lower Pleistocene: Atlantic, endemic to Selsoif, NW France (this paper).
Shell small, turritelliform. Protoconch paucispiral, consisting of one convex whorl with raised nucleus. Teleoconch of at least seven strongly convex whorls separated by moderately impressed linear suture. Sculpture on first teleoconch whorl of three elevated cords, adapical cord placed on subsutural ramp weakest, middle cord delimiting shoulder strongest, abapical cord slightly weaker than mid-cord. Abapically, upper cord strengthens, so that on third teleoconch whorl three strong primary cords, mid-cord still slightly stronger forming whorl periphery, fourth cord appears on subsutural ramp. On fourth whorl a fifth primary cord appears on subsutural ramp; later whorls with five primary spiral cords, upper two slightly weaker than lower three. Fine, elevated axial lamellae present in spiral interspaces. Last whorl short, strongly convex, with six cords above the level of the insertion of the outer lip plus weak peribasal cord; base smooth. Aperture small, rounded, outer lip simple, oblique in profile.
The late Neogene of NW France is proving fertile hunting ground for endemic et al. 2018) . et al. (2018) have the more usual number of three spiral cords on all adult whorls. It is also separated from all the present-day Atlantic species discussed by Rolán & Fernandes (1990) that have multispiral protoconchs. It is also unusual in having strongly convex whorls. Most
-
(Fig. 6 E1-E3, F1-F3, G1-G3)
Height 4.6 mm (incomplete), width 1.7 mm. — RGM.1365212 (1), leg. ACJ; RGM.1365310 (9), leg. ACJ; RGM.1365307 (1), leg. ACJ; RGM.1365306 (1), leg. ACJ; RGM.1365308 (2), leg. ACJ; RGM.1364977 (23), leg. ACJ; RGM.1365309 (1), leg. ACJ; RGM.1365311 (1), leg. ACJ; RGM. 1365313 (1) leg. unknown, RGM.1365078 (c. 135) leg AWJ, RGM.961903 (26), leg. AWJ; RGM.961908 (7), leg. unknown.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
-
-
(Fig. 7 A1-A3, B1-B3)
nonnon J. de C. Sowerby, 1813).
partim): 90, pl. 8, fig. 11 (fossil specimens only)[ non
et al. 2012 : 174, fig. 18.
Maximum height 35.4 mm, width 13.7 mm. — RGM.1364910 (24), leg. WG; RGM.1365181 (1), leg. WG; RGM.1364911 (1), leg. WG; RGM.1364979 (34), ACJ.
Robust epitoniid with variable convex to subconvex whorl profile and a characteristic spiral ornamentation of low broad ribs. The whorl profile, number and strength of the ribs, including a basal rib, are very variable.
Lower Pliocene: NSB, Coralline Crag Formation, England (Harmer 1925). — Upper Pliocene: NSB, Red Crag Formation, England (Wood 1848; Harmer 1925), Netherlands ( Wesselingh et al. 2012) . — Lower Pleistocene: NSB, England (Harmer 1925); Atlantic, Selsoif, NW France (this paper).
et al. (2012 : compare fig. 17 vs fig. 18).
It is tempting to suggest that et al. (2021 : pl. 7.8, fig. 6) from the Late Pliocene-Early Pleistocene of Iceland is clearly that species with its shallow suture and weakly convex whorls. Durham & MacNeil (1967) suggested that
It is also possible that the two represent extreme forms of a single species, and that the differences seen are ecophenotypic. Despite the variability seen in both species, Pliocene specimens in the RGM collections could relatively easily be ascribed to
-
-
(Fig. 7 C1-C3)
Maximum height 10.2 mm (incomplete), width 6.3 mm. — RGM.1365195 (1), leg. WG; RGM.1364912 (2), leg. WG; RGM.1365173 (1), leg. ACJ; RGM.1365096 (17), leg. AWJ.
Smooth, medium-sized shell. Protoconch damaged or lacking in almost all specimens, but in a few specimens the first whorl is flat and not clearly separated from the teleoconch. Teleoconch of four round whorls. Aperture typically circa 40% of shell height. Umbilicus reduced to narrow slit.
Lower Pleistocene: NSB, England (Harmer 1921); Atlantic, Selsoif, NW France (this paper). — Holocene: England (Harmer 1921), West Greenland (Símonarson 1981). Beached occurrences of this species likely of a Late Pleistocene age are known from The Netherlands (Moerdijk & Janse 2015). Today this species occurs from Greenland, Arctic Canada to the British Isles and Normandy (Fretter & Graham 1980; Backeljau et al. 1984) . The species has been reported living on or near the fleshy bryozoan
(Fig. 7 D1-D3)
Maximum height 4.7 mm, width 5.1 mm. RGM.1364981 (20), leg. ACJ; RGM.1365182 (1), leg. AWJ; RGM.1364913 (2), leg. WG; RGM.1365081 (51), leg. AWJ.
Small, round shell. Three smooth teleoconch whorls; spire almost flat; last whorl 92% of the total height. Prosocline growthlines. Aperture round; moderately extent columella fold on an open umbilicus.
Lower Pleistocene: Scotland and Ireland (Harmer 1921); Atlantic, Selsoif, NW France (this paper). — Holocene: NSB (Strand Petersen 2004). Today this is predominantly a northern species extending from south of the Arctic Circle on the American coast from Greenland to Connecticut (Gould 1840; Trott 2004), in the eastern Atlantic the distribution extends from the White Sea ( Vortsepneva et al. 2023) , North Sea (Dumoulin 1989), British Isles (Fretter & Graham 1980; Guiry & Guiry 2011), France (Müller 2004) into northern Spain (Rolán 1983). The species is associated with
(Fig. 7 E1-E3)
Maximum height 3.5 mm, width 3.4 mm RGM.1364980 (4), leg. ACJ; RGM.1365183 (1), leg. ACJ.
Small ovate shell. Three smooth, round teleoconch whorls. Last whorl is 77% of total shell. Aperture round with a thick peristome, columella expanded and flattened abapically. Broad umbilical chink.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). — Lower and Upper Pleistocene: England and Ireland (Harmer 1921). — Upper Pleistocene: Netherlands (Wolf 2001). — Holocene: NSB (Strand Petersen 2004). Today predominantly a northern species extending from south of the Arctic Circle on the American coast to Cape Cod, in the eastern Atlantic British Isles, Norway, Sweden (Fretter & Graham 1980), North Sea of Belgium (Backeljau 1986). The species is associated with algae such as
(Fig. 7 F1-F3)
cf.
cf.
Maximum height 2.7 mm (incomplete), width 2.1 mm. — RGM.1364982 (1), leg. ACJ; RGM.1365184 (2), leg. ACJ.
Small, ovate-conical shell with subobsolete spiral threads. Four teleoconch flat whorls. Last whorl is 59% of the total height. Prosocline growth-lines. Ovate aperture, with open umbilicus.
Selsoif, NW France (this paper). Today
We have only badly damaged material, whose attribution is uncertain. Nevertheless, the flat-sided whorl profile and subobsolete spiral threads suggest it concerns this species.
-
(Fig. 7 G1-G3, H1-H3)
partim): 661, pl. 52, figs 27-28 (non fig. 29, that corresponds to
For more, see synonymy list in Reid (1996).
Maximum height 8.4 mm, width 9.3 mm. RGM.1365211 (1), leg. ACJ; RGM.1364983 (6), leg. ACJ; RGM.1365076 (7), leg. WG., RGM.1310829 (1), leg. WG; RGM.1365087 (3), leg AWJ, RGM. 1365286 (24) leg. WG; RGM.1365086 (c. 90) leg. AWJ; RGM.1405527 (46), leg ACJ.
Characterisation. — Relatively small (H = 6 mm) thick shelled, robust, often smooth and shiny viz.Reid 1996); aperture rounded with particularly thickened abapical margin; well preserved specimens bear very fine, slightly irregular spiral grooves. The ratio between the height of the lower apertural lip and the height of the outerlip ranges between 0.20 and 0.29, which is within the range of
The published fossil distribution of
Today in western Europe two closely resembling viz. the ratio between basal lip height versus the aperture height. The dated phylogenies of Reid (1996) indicate they are sister species that evolved from a common ancestor around 2 Ma, broadly contemporary with the Selsoif material.
(Fig. 7 I1-I3, J1-J3)
et al. 1954 : 61, fig. 29. — Moerdijk & Janse 2015: 20, figs 1-2.
nonet al. 1980 (partim): 65, pl. 13, figs 8-9.
nonet al. 1980 (partim), p. 65, pl. 13, figs 10-11 (that corresponds to
Maximum height 25.3 mm (incomplete), width 22.6 mm. RGM.1365202 (100+) leg. AWJ; RGM.1365070 (1), leg. AWJ; RGM.1364984 (37), leg. ACJ; RGM.1365088 (82), leg. AWJ; RGM.1365086 (56), leg. WG; RGM.1364914 (28), WG.
Globular to ellipsoid, thick shell. Short spire with a pointed apex. Juveniles have strong spiral ribs that become obsolete early on the teleoconch. On some specimens the last whorl has at least three brown or red spiral lines. Even though the strength and location of disappearance of spiral ribs is somewhat variable, the overall shape is very distinct.
Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1920; Reid 1996). — Lower Pleistocene: NSB, England (Harmer 1920), Netherlands ( Van Regteren Altena et al. 1954 ; Moerdijk & Janse 2015); Atlantic, Iceland (not in Icelandic modern fauna) (Reid 1996), Selsoif, NW France (this paper). — Middle Pleistocene: Channel region England (Preece & Bates 1999). — Upper Pleistocene: NSB, England (Harmer 1920). Recent distribution in the western Atlantic from Newfoundland to Chesapeake Bay in the western Atlantic (Reid 1996), in the eastern Atlantic from the White Sea southwards, occasionally as far as southern Portugal. The lack of fossil occurrences in the western Atlantic allows the possibility that the species was introduced there by e.g., Vikings (Reid 1996).
-
-
(Fig. 8 A1-A3)
cf.
Maximum height 2.6 mm, width 1.6 mm. — RGM.1364986 (1), leg. ACJ; RGM.1310338 (1), leg ACJ.
Shell small, rissoiform. Protoconch not preserved. Three teleoconch whorls preserved. Sculpture of three spiral cords on spire whorls, and 13 orthocline ribs forming coarsely reticulated sculpture. Last whorl bearing six spiral cords. Aperture round, peristome complete, thickened, outer lip thickened.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). et al. (2013) reported occasional findings from the Upper Pleistocene NSB of the Netherlands. Today it is distributed from the British Isles south into the Mediterranean ( Landau et al. 2004) .
Due to the scarcity and poor preservation of the material at hand, identification is uncertain. It is clearly separated from the other two congeners present in the assemblage by the shape and the number of spiral cords.
(Fig. 8 B1-B3)
urn:lsid:zoobank.org:act:19AACEFD-F1BE-4FF6-B380-411C58ACC056
RGM.1365286, height 1.4 mm, width 0.9 mm, leg. ACJ.
Named after the type locality of Selsoif.
Selsoif, Saint-Sauveur-le-Vicomte, Manche department, NW France.
Gelasian, Early Pleistocene.
Lower Pleistocene: Atlantic, endemic to Selsoif, NW France (this paper).
Shell small, rissoiform. Protoconch of two smooth whorls (surface sculpture may be abraded). Three teleoconch whorls; sculpture on spire whorls of two elevated spiral cords; axial sculpture of 12-13 narrow, elevated, widely spaced, orthocline ribs forming coarsely reticulated pattern. Whorl profile sharply angled at the cords. Last whorl 60% of total height, single secondary develops on second half of last whorl, profile sharply angular at shoulder and peripheral cord, peribasal cord prominent with two further cords of equal strength over base, axials weakening over base: base somewhat depressed. Aperture 35% of total height, ovate; outer lip thickened by varix, peristome complete; columella thickened in abapical portion.
If we review species with similar shell characters outside the Mediterranean,
(Fig. 8 C1-C3)
Maximum height 1.3 mm, width 0.8 mm. — RGM.1364985 (8), leg. ACJ; RGM.1365091 (107), leg. AWJ; RGM.1365090 (1), leg. AWJ; RGM.1365089 (10), leg. AWJ; RGM.1364987 (2), leg. ACJ.
Shell small, rissoiform. Protoconch of two smooth whorls. Three teleoconch whorls; spire whorls sculpture of four spiral cords; axial sculpture of around 16 broad orthocline ribs forming a reticulated pattern. Last whorl develops one secondary cord between suture and first cords. Base somewhat depressed, with four spiral cords; axial ribs stop at peribasal cord.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
The material at hand is too worn to identify with certainty. In profile and sculpture, it is similar to several Italian Pliocene species such as
-
(Fig. 8 D1-D3)
et al. 2016 : 140, pl. 10, fig. 2. — Landau et al. 2018 : 274, pl. 103, fig. 1.
For more, see synonymy list in Van Dingenen et al. (2016) and Landau et al. (2018) .
Maximum height 1.7 mm, width 1.0 mm. RGM.1364988 (4), leg. ACJ; RGM.1365201 (1), leg. ACJ; RGM.1365093 (16), leg. AWJ.
Shell small, rissoiform. Protoconch not preserved. Four strongly convex teleoconch whorls separated by deeply impressed suture; sculpture of numerous, regular, flattened spiral cords; about ten high, widely spaced, sinuous, prosocline axial ribs. Ribs stop abruptly at strong, elevated peribasal cord; base narrow, deeply concave forming wide groove between peribasal cord and peristome. Aperture ovate, outer lip strongly thickened by rounded labial varix, peristome complete.
Middle Miocene: Atlantic, Loire Basin, France (de Morgan 1915; Cossmann 1918; Glibert 1949); Paratethys, Austria (Kowalke & Harzhauser 2004), Hungary (Strausz 1966), Poland (Friedberg 1923; Bałuk 1975), Romania (Kowalke & Harzhauser 2004); Proto-Mediterranean, Karaman Basin, Turkey ( Landau et al. 2013) . — Upper Miocene: Atlantic, NW France (Calas 1949; Landau et al. 2018) ; Proto-Mediterranean, Italy (Sacco 1895; Venzo & Pelosio 1963). — Lower Pliocene: Atlantic of NW France ( Van Dingenen et al. 2016) ; central Mediterranean, Italy (Chirli 2006), Tunisia (Fekih 1975). — Upper Pliocene: Atlantic, Portugal (Silva 2001); western Mediterranean, Spain ( Landau et al. 2004) ; central Mediterranean, Italy (Sacco 1895; Malatesta 1974; Cavallo & Repetto 1992; Sosso & dell’Angelo 2010; Tabanelli et al. 2011) . — Lower Pleistocene: NSB, British Isles (Harmer 1918); Atlantic, Selsoif, NW France (this paper); western Mediterranean of France (Glibert 1962); central Mediterranean, Italy (Cerulli-Irelli 1914; Buccheri 1970); eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Upper Pleistocene: NSB, the Netherlands ( Van Regteren Altena et al. 1954 ; Raad et al. 2013) ; western Mediterranean, Balearic Islands (Cuerda Barceló 1987). Today this species occurs in the northeastern Atlantic from Norway southwards to Morocco, entire Mediterranean, from extreme low tide line to 50 m depth, often on sandy bottoms, but also under stones and in weeds (Poppe & Goto 1991).
For discussion on fossil populations, see Landau et al. (2013 : 74) and Van Dingenen et al. (2016 : 140).
-
(Fig. 8 E1-E3)
et al. 1954 : 12, pl. 4, fig. 35b. — A.W. Janssen 1967: 128, pl. 4, fig. 7.
et al. 2013 : 52, fig. 19. — Símonarson & Eiríksson inEiriksson & Símonarson 2021: 264, pl. 7.3, fig. 5.
Maximum height 1.9 mm, width 0.8 mm. — RGM.1364989 (27), leg. ACJ; RGM.1365197 (1), leg. ACJ; RGM.1365095 (800+), leg. AWJ.
Shell small, rissoiform. Smooth protoconch of 1.5 whorl; five round teleoconch whorls separated by deep suture. Adult sculpture consist of axial ribs marked on the upper whorls and numerous fine crowded spiral cords covering the entire whorl. Ovate aperture; rounded, thickened outer lip.
Middle Miocene: NSB, Germany (Janssen 1967; Wienrich 2001). — Lower Pliocene: Coralline Crag, England (Wood 1848; Harmer 1920) and Belgium (Marquet 1997). — Upper Pliocene: NSB, Belgium (Marquet 1997). — Pliocene to Lower Pleistocene: Iceland (Eiriksson & Símonarson 2021). — Lower Pleistocene: Atlantic, England (Harmer 1920; Norton 1967); Selsoif, NW France (this paper). — Pliocene and Pleistocene (indeterminate): NSB, Netherlands ( Van Regteren Altena et al. 1954 ; Raad et al. 2013) . — Holocene: NSB (Strand Petersen 2004). Today this species is present in the Atlantic from Iceland (Warén 1996), Norway (Høisæter 2009), Kola Peninsula, Russia (Golikov & Kussakin 1978; Nekhaev et al. 2014) , British Isles (Fretter & Graham 1978), Madeira ( Ávila et al. 2012) , southwards to the Alboran Sea in the Mediterranean (Templado & Rolán 1986). Shells collected in the central Mediterranean are probably Late Pleistocene glacial fossils (Amati & Nofroni 2015). Abundant under rocks and among algae, from the intertidal to 1000 m depth.
-
(Fig. 8 F1-F3)
Maximum height 1.3 mm, width 0.8 mm. — RGM.1365258 (1), leg. ACJ; RGM.1365125 (5), leg. AWJ; RGM.1365097 (62), leg. AWJ; RGM.1405528 (2), leg. ACJ; RGM.794506 (1), leg. AWJ.
Small rather high-spired rissoiform shell. Protoconch not preserved. Five convex teleoconch whorls separated by a deep suture. Sculpture variable mostly smooth but with some specimens consisting of numerous sinuous, opisthocline ribs strengthening towards abapical suture. Base rounded, smooth. Aperture ovate, angled at anal sinus.
The overall characters resemble those of
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
-
(Fig. 9 A1-A3)
et al. 2013 : 55, fig. 26.
non Montagu, 1803). — Harmer 1920: 634, pl. 51, fig. 16 (not fig. 17).
et al. 1954 : 13, pl. 4, fig. 41. — Marquet 1997: 15, pl. 2, fig. 4.
Maximum height 1.6 mm, width 1.0 mm. — RGM.1364990 (273), leg. ACJ; RGM.1365196 (1), leg. AWJ; RGM.1365092 (66), leg. AWJ; RGM.1364915 (4), leg. WG.
Small broad rissoiform shell. Three, flattened protoconch whorls. Four teleoconch whorls. Sculpture of around 16 strong, orthocline ribs covering numerous spiral ribs. Base covered by five weak spiral cords. Umbilicus closed; aperture ovate, thick, denticulate outer lip.
Upper Pliocene: NSB, Red Crag, England (Harmer 1920), Belgium (Marquet 1997). — Pliocene to Lower Pleistocene (indeterminate): Netherlands ( Van Regteren Altena et al. 1954 ; Raad et al. 2013) . — Lower Pleistocene: NSB, England (Wood 1848; Harmer 1920); Atlantic, Selsoif, NW France (this paper).
Wood (1848: 102) figured this species under the name
(Fig. 9 B1-B3)
et al. 1998 : 31, figs 12d, 15. — Raad et al. 2013 : 57, figs 31-32.
et al. 1954 : 13, pl. 4, fig. 42.
Maximum height 2.4 mm, width 1.1 mm. — RGM.1364991 (89), leg. ACJ; RGM.1365260 (1), leg. ACJ; RGM.1365035 (7), leg. AWJ; RGM.1365094 (60), leg. AWJ.
Small elongated rissoiform shell. Up to six convex teleoconch whorls. Sculpture only on the three last whorls of around eight axial ribs separated by wider interspace. Base smooth. Aperture pyriform; columella oblique; imperforate.
Lower Pliocene: NSB, Coralline Crag, England (Harmer 1920). — Lower and Upper Pleistocene: British Isles (Harmer 1920). — Lower Pleistocene: Atlantic, Greenland ( Símonarson et al. 1998 ), Selsoif, NW France (this paper). — Upper Pleistocene to Holocene: Netherlands ( Van Regteren Altena et al. 1954 ; Raad et al. 2013) . — Holocene: NSB (Harmer 1920; Strand Petersen 2004). Today, this species is present along the entire European Atlantic coast, Norway ( Christie et al. 2003) , North Sea (Ziegelmeier 1966; de Bruyne 1991), entire Mediterranean and into the Aegean ( Öztürk et al. 2014) and Black Sea ( Öztürk et al. 2014) .
This species shows large morphological variations in the shape, size and whorl profile, as well as ornamentation (from smooth to strongly ribbed on the three last whorls).
-
-
(Fig. 9 C1-C3)
cf.
cf.?
cf.
cf.
cf. inEiriksson & Símonarson 2021: 256, pl. 7.2, fig. 8.
Maximum height 1.5 mm, width 0.8 mm. — RGM.1365261 (1), leg. ACJ.
Thin shelled hydrobiid with marked convex whorl profile and relatively deep suture; aperture rounded. The specimen at hand suffers from corrosion of the outer shell layer, hence its uncertain attribution.
Lower Pliocene: Iceland ( Símonarson et al. 2021 ). — Upper Pliocene: NSB, Red Crag, England (Hoeksema & Raad 2015). — Lower Pleistocene: Maassluis Formation, Netherlands (RGM collection); Atlantic, Selsoif, NW France (this paper). — Pleistocene and Holocene (indeterminate): NSB, Netherlands (Hoeksema & Raad 2015). Today it occurs along the European coasts from Norway and the Baltic to the Mediterranean and Black Sea (Fretter & Graham 1978).
-
(Fig. 9 D1-D3)
et al. 1954 : 12, pl. 3, fig. 32. — Símonarson & Eiríksson inEiriksson & Símonarson 2021: 255, pl. 7.2, fig. 7.
Maximum height 2.1 mm, width 1.1 mm. — RGM.1364993 (10), leg. ACJ; RGM.1365198 (1), leg. ACJ; RGM.1365202 (200+), leg. AWJ.
Smooth, solid, fusiform shell with regularly conical spire. Five flat teleoconch whorls separated by shallow suture. High base; pyriform aperture.
Lower Pliocene: NSB, Coralline Crag, England (Wood 1848), Iceland ( Símonarson et al. 2021 ). — Upper Pliocene: NSB, Red Crag, England (Harmer 1925; Norton 1967). — Lower Pleistocene: Atlantic, England (Harmer 1925), Selsoif, NW France (this paper). — Upper Pleistocene to Holocene: NSB, England (Harmer 1925), Netherlands ( Van Regteren Altena et al. 1954 ; Hoeksema & Raad 2015). Today, this species occurs in the eastern Atlantic from northern Norway (Bank & Neubert 2020), along the coast of Europe to Senegal (Fretter & Graham 1978), it enters the Mediterranean and extends into the Aegean Sea ( Öztürk et al. 2014 ) and Baltic Sea (Kedra 2010).
-
-
(Fig. 9 E)
et al. 2016 : pl. 11, fig. 5.
For more, see synonymy list in Van Dingenen et al. (2016) .
Maximum length 1.1 mm. — RGM.1365099 (52), leg. AWJ; RGM.1365191 (1), leg. AWJ.
Small, elongate, smooth shell. Dome-shaped apical septum.
Middle Miocene: Loire Basin, France (Glibert 1949). — Lower Pliocene: Atlantic, NW France ( Van Dingenen et al. 2016) ; NSB, Coralline Crag, England (Wood 1848; Harmer 1923). — Upper Pliocene: NSB, Red Crag, England (Harmer 1923); Oorderen Sands, Belgium (Marquet 1997). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper); central Mediterranean, Italy (Cerulli-Irelli 1912). — Pleistocene (indeterminate): Atlantic, British Isles (Harmer 1923); NSB, The Netherlands ( Van Regteren Altena et al. 1955 ). — Holocene: NSB (Strand Petersen 2004). Today this species occurs at the Atlantic coasts of Europe from British Isles to the Bay of Biscay (Fretter & Graham 1978). Mediterranean records are not
-
-
(Fig. 9 F1-F3)
et al. 2016 : 149, pl. 11, fig. 9, pl. 12, fig. 3. — Landau et al. 2018 : 303, pl. 131, fig. 1.
For more, see synonymy list in Van Dingenen et al. (2016) and Landau et al. (2018) .
Maximum height 0.8 mm, width 1.2 mm. — RGM.1364994 (19), leg. ACJ; RGM.1365009 (93), leg. AWJ; RGM.1365299 (1), leg. AWJ; RGM.1405531 (1), leg. ACJ.
Small, depressed, subcircular shell. Protoconch of 2.5 smooth whorls; three teleoconch whorls with carinate shoulder, mid-whorl and peribasal cords, sinuous ribs on dorsum and venter; base flattened, bearing two further elevated cords, the medial periumbilical cord delimiting wide, deep umbilicus with ribs extending within; aperture wide, rounded, strongly oblique in profile.
Middle Miocene: Atlantic, Loire Basin, France (Glibert 1949). — Upper Miocene: Atlantic, NW France ( Landau et al. 2018) . — Lower Pliocene: NSB, Coralline Crag, England (Wood 1848; Harmer 1923; Atlantic, NW France ( Van Dingenen et al. 2016) ; central Mediterranean, Italy (Chirli 2006). — Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1923). — Upper Pliocene-Pleistocene: Atlantic, NW France (Cossmann 1918; Brébion 1964). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). — Upper Pleistocene: NSB, The Netherlands ( Van Regteren Altena et al. 1954 ); Atlantic, British Isles (Harmer 1923). Today this species is present in the Atlantic coasts of Europe from British Isles to Tarifa (Fretter & Graham 1978).
Van Dingenen et al. (2016 :149) highlighted the problem with the et al. 1998 : fig. 7), and a Mediterranean species that is at present unnamed with a paucispiral protoconch of 1.25-1.4 whorls, with a larger nucleus (Rolán & Rubio 2002: figs 13, 14; Van Aartsen et al. 1998 : fig. 7). With a protoconch of 2.5 whorls the specimens from Selsoif are typical of the Atlantic species
-
-
(Fig. 10 A)
et al. 2013 : 95, pl. 9, fig. 7, pl. 61, fig. 6. — Chirli & Forli 2017: 52, fig. 15a. — Landau et al. 2018 : 316, pl. 147, fig. 1. — Sacchetti et al. 2023 : 51, pl. 3, fig. F.
inEiríksson & Símonarson 2021: 267, pl. 7.4, fig. 1.
For more, see synonymy list in Landau et al. (2013 , 2018).
Maximum height 4.8 mm, diameter 11.4 mm. — RGM.1364916 (2), leg. WG; RGM.1364995 (127), leg. ACJ; RGM.1365194 (1), leg. ACJ; RGM.1365100 (2), leg. AWJ; RGM.1405530 (18), leg. WG.
Relatively large, smooth, strongly depressed conical shell. Apex subcentral, consisting of only protoconch whorl. Aperture entire venter, septum with sinuous border extending from mid-shell to outer edge.
Lower Miocene: Paratethys, Austria (Schaffer 1912); Proto-Mediterranean: Italy (Sacco 1896b). — Lower-Middle Miocene: NSB: Belgium (Glibert 1952), Germany (Anderson 1964), the Netherlands (Janssen 1984). — Middle Miocene: Atlantic, Aquitaine Basin, France (Cossmann & Peyrot 1919), Loire Basin, France (Glibert 1949); Paratethys, Poland (Friedberg 1923; Bałuk 1975), Austria (Hornes 1856; Schultz 1998), Bulgaria (Kojumdgieva & Strachimirov 1960), Hungary (Strausz 1966), Ukraine ( Zelinskaya et al. 1968) , eastern Paratethys (Iljina 1993); Proto-Mediterranean, NE Spain (Solsona 1998), Karaman Basin, Turkey (Fischer 1866; Landau et al. 2013) . — Upper Miocene: Atlantic, NW France ( Landau et al. 2020a) , Portugal (Glibert 1963), SW Spain ( Cárdenas et al. 2019) ; Proto-Mediterranean, Italy (Sacco 1896b; Venzo & Pelosio 1963; Boschele et al. 2021) . — Lower Pliocene: NSB, Coralline Crag, England (Wood 1848; Harmer 1921), Belgium (Glibert 1958; Marquet 1998), Iceland (Eiríksson & Símonarson 2021), Atlantic, Santa Maria Island, Azores ( Sacchetti et al. 2023) ; NW France ( Van Dingenen et al. 2016) , SW Spain (Gonzalez-Delgado 1988; Landau et al. 2011 ; Brunetti 2022), Morocco (Lecointre 1952); NE Spain (Martinell 1979), Roussillon Basin, France (Fontannes 1879); central Mediterranean, Italy (Sacco 1896b; Palla 1967; Caprotti 1974; Brunetti & Cresti 2018), Tunisia (Fekih 1975). — Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1921), Belgium (Glibert 1958; Marquet 1998); Atlantic, Pombal Basin, Portugal (Silva 2001), Morocco (Lecointre 1952); western Mediterranean, S. Spain ( Landau et al. 2004) ; central Mediterranean, Italy (Malatesta 1974; Chirli 1988; Cavallo & Repetto 1992). — Upper Pliocene-Pleistocene: NW France (Brebion 1964). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). — Pleistocene: Atlantic, British Isles (Glibert 1963), Morocco (Lecointre 1952); central Mediterranean, Italy (Cerulli-Irelli 1912; Taviani et al. 1998 ; Brunetti & Vecchi 2015; Chirli & Forli 2017), Sicily (Glibert 1963); eastern Mediterranean (Chirli & Linse 2011). Today this species occurs in the NE Atlantic, British Isles to Zaire, Madeira and Canaries, Mediterranean and Black Sea (Poppe & Goto 1991).
As discussed by Landau et al. (2004 , 2013, 2020a), this species has an extremely variable shell in both profile profile; tall conical to almost flat, and sculpture; smooth or bearing small spines. The size and position of the protoconch varies between populations and it is possible that it represents a species complex rather than a single taxon (B. Landau & R. Pouwer pers. obs.).
-
(Fig. 10 B1, B2)
et al. 2018 : 317, pl. 148, fig. 1.
Maximum height 3.5 mm, diameter 12.3 mm. — RGM.1364917 (1), leg. WG; RGM.1364996 (2), leg. ACJ; RGM.1365101 (62), leg. AWJ; RGM.1365185 (1), leg. AWJ.
Thick, low, smooth, convex shell. Aperture ovate. Concave, shallow septum covering almost 2/3 of the aperture.
Middle Miocene: Atlantic, Aquitaine Basin, France (Cossmann & Peyrot 1919), Loire Basin, France (Glibert 1949); Paratethys, Austria (Hörnes 1856), Poland (Bałuk 1975), Hungary (Strausz 1966); Proto-Mediterranean, Karaman Basin, Turkey ( Landau et al. 2013) . — Upper Miocene: Atlantic, NW France ( Landau et al. 2018 ); Proto-Mediterranean, Italy (Sacco 1896b). — Lower Pliocene: Atlantic, NW France ( Van Dingenen et al. 2016) ; SW Spain (González-Delgado 1988; Landau et al. 2011) . — Upper Pliocene: Atlantic, Morocco (Lecointre 1952); western Mediterranean, S Spain ( Landau et al. 2004) ; central Mediterranean, Italy (Chirli 1988; Cavallo & Repetto 1992). — Upper Pliocene-Pleistocene: NW France (Glibert 1963). — Lower Pleistocene, Atlantic, Selsoif, NW France (this paper); central Mediterranean (Chirli & Forli 2017). Today, this species occurs in the Mediterranean Sea, below low tide line, often attached to other molluscs (Poppe & Goto 1991).
Our assignment of et al. (2004 : 74). In our opinion the taxonomy of European
-
-
(Fig. 10 C1, C2)
sic ] – Domènech & Martinell 1982: 50, fig. 15.
et al. 2011 : 14, pl. 4, fig. 7. — Chirli & Linse 2011: 63, pl. 14, fig. 5. — Van Dingenen et al. 2016 : 123, pl. 4, fig. 4. — Chirli & Forli 2017: 54, fig. 15d.
For more, see synonymy list in Landau et al. (2011) and Van Dingenen et al. (2016) .
Maximum height 25.2 mm, width 11.9 mm (incomplete). — RGM.1364918 (4), leg. WG; RGM.1364997 (18), leg. ACJ; RGM.1365102 (57), leg. AWJ; RGM.1365200 (1), leg. AWJ; RGM.1364998 (1), leg. ACJ.
Medium sized capuliform shell with ovate aperture, convex elevated in profile with coiled, recurved apex displaced slightly to the right overhanging posterior margin. Protoconch of just under two smooth whorls with small nucleus, rapidly expanding last whorl sculpture of fine axial ribs. U-shaped internal muscle scar.
Middle Miocene: Paratethys, Poland (Bałuk 1975), Vienna (Hörnes 1856); Proto-Mediterranean, Italy (Sacco 1896b); Atlantic, Aquitaine Basin, France (Cossmann & Peyrot 1919), Loire Basin, France (Glibert 1949). — Upper Miocene: Proto-Mediterranean, Italy (Sacco 1896b). — Lower Pliocene: NSB, Coralline Crag, England (Wood 1848; Harmer 1916), Kattendijk Formation, Belgium (Glibert 1958; Marquet 1998); Atlantic, NW France ( Van Dingenen et al. 2016) , SW Spain ( Landau et al. 2011) ; central Mediterranean, Italy (Chirli 2008). — Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1916), Oorderen Sands, Belgium (Glibert 1958; Marquet 1998); Atlantic, Portugal (Zbyszewski 1959; Silva 2001); western Mediterranean, S Spain ( Landau et al. 2004) , France (Chirli & Richard 2008); central Mediterranean, Italy (Sacco 1896b; Cavallo & Repetto 1992). — Upper Pliocene-Pleistocene: NW France (Brébion 1964). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper); central Mediterranean, Italy (Cerulli-Irelli 1914; Brambilla & Galli 1988; Di Geronimo & La Perna 1997; Chirli & Forli 2017); eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Upper Pleistocene: western Mediterranean, Spain (Domènech & Martinell 1982). Today this species occurs in the Atlantic from Iceland to West Africa and North America to Florida, Mediterranean. It is not present in the North Sea (Fretter & Graham 1981). It inhabits in the sublittoral zone to 850 m depth, attached to stones and shells (Poppe & Goto 1991)
The Selsoif specimens have a protoconch of about two whorls, similar to that illustrated by Fretter & Graham (1981: fig. 225). For further discussion see Van Dingenen et al. (2016) .
-
-
(Fig. 10 D1-D3)
et al. 1955 : 33, fig. 77. — Menkhorst & Wesselingh 2018: 42, figs 13, 14. — Landau & Mulder 2022: 161, figs 10a-c.
For more, see synonymy list in Landau & Mulder (2022).
Maximum height 1.8 mm (incomplete), width 0.6 mm. — RGM.1365077 (11), leg. AWJ; RGM.1365257 (1), leg. AWJ.
Small, turriform shell with narrow apical angle. Protoconch absent in all specimens. About six convex teleoconch whorls separated by narrowly impressed suture. Three spiral cords on early whorls, a fourth one appearing above the suture on later whorls; aperture ovate.
Upper Miocene: NSB, Denmark (Rasmussen 1968; Schnetler 2005), Germany (Moths 1989). — Lower Pliocene: NSB, Coralline Crag, England (Wood 1842, 1848; Harmer 1925); central Mediterranean, Italy (Chirli 2009; Brunetti & Cresti 2018). — Upper Pliocene: western Mediterranean, S Spain ( Landau et al. 2006 ; Landau & Mulder 2022). — Lower Pleistocene: NSB, England, Ireland (Harmer 1925); Atlantic, Selsoif, NW France (this paper); central Mediterranean, Italy (Cerulli-Irelli 1914); eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Pleistocene (indeterminate): NSB, Netherlands ( Van Regteren Altena et al. 1955 ; Menkhorst & Wesselingh 2018). — Holocene: NSB (Strand Petersen 2004). Today this species is present in the Atlantic, from the British Isles (Guiry & Guiry 2011) to Madeira ( Segers et al. 2009) and in the Mediterranean Sea from Spain ( Peñas et al. 2009) to the Aegean Sea ( Öztürk et al. 2014) , at depths of 2-250 m, on calcareous algae, sandy, or muddy bottoms ( Albanesi et al. 1981 ).
(Fig. 10 E1-E3)
et al. 1955 : 21, fig. 78. — Menkhorst & Wesselingh 2018: 43, fig. 15.
non Aclis ascaris (Turton, 1819)].
Maximum height 2.4 mm, width 1.1 mm. — RGM.1365001 (1), leg. ACJ.
Small, smooth, turriform shell with relatively broad apical angle. Protoconch missing. Up to six low, convex teleoconch whorls separated by deep suture; smooth, somewhat flattened base; aperture incomplete.
Lower Pliocene: NSB, Coralline Crag, England (Wood 1848; Harmer 1925). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). — Upper Pleistocene: NSB, Netherlands ( Van Regteren Altena et al. 1955 ; Menkhorst & Wesselingh 2018). Today this species occurs from Iceland to Spain (Bouchet & Warén 1986), Azores (Hoffman & Freiwald 2020).
-
(Fig. 10 F1-F3)
et al. 2018 : 313, pl. 143, figs 1-3. — Menkhorst & Wesselingh 2018: 38, fig. 1.
For more, see synonymy list in Landau et al. (2018) .
Maximum height 11.7 mm, width 3.7 mm. — RGM.1364919 (112), leg. WG; RGM.1365000 (67), leg. ACJ; RGM.1365103 (c. 50), leg. AWJ; RGM.1365318 (1), leg. AWJ.
Relatively large, slender, elongate shell, slightly curved with smooth polished surface without magnification, finely reticulated sculpture under magnification. Nine flat teleoconch whorls separated by a shallow suture. Base flat; columellar lip short, thickened, almost straight to inclined; aperture pyriform.
Middle Miocene: NSB, Netherlands (Janssen 1984). — Upper Miocene: Atlantic, NW France ( Landau et al. 2018) . — Lower Pliocene: Atlantic, NW France ( Van Dingenen et al. 2016) ; NSB, Coralline Crag, England (Harmer 1920), Kattendijk Formation, Belgium (Marquet 1998). — Upper Pliocene: NSB, Red Crag, England (Harmer 1920), Atlantic, Portugal ( Landau et al. 2006) ; western Mediterranean, S Spain ( Landau et al. 2006) . — Pliocene (unspecified): NSB, the Netherlands ( Van Regteren Altena et al. 1955 ; Menkhorst & Wesselingh 2018); Atlantic, St Erth, England (Harmer 1920), Selsoif NW France (this paper). — Holocene: NSB (Strand Petersen 2004). Today this species is present in the Atlantic, from Norway and rare into the Mediterranean (Fretter & Graham 1982).
-
-
(Fig. 10 G1-G3)
aff.
aff.
aff. et al. 1965 : 26, pl. 10 fig. 99. — Símonarson & Eiríksson inEiríksson & Símonarson 2021: 269, pl. 7.4, fig. 2. — Van Nieulande et al. 2022 : 16, fig. 1.
Maximum height 3.5 mm, width 4.5 mm. — RGM.1364920 (1), leg. WG; RGM.1365002 (4), leg. ACJ; RGM.1365186 (1), leg. AWJ; RGM.1365104 (93), leg. AWJ; RGM.1365206 (2), leg. AWJ.
Capuliform shell with paucispiral protoconch and rapidly expanding last whorl resulting in large circular aperture, disjunct last whorl.
Lower Pleistocene: Atlantic, Selsoif, France (this paper).
The specimens from Selsoif differ from
Disjunct last whorls occur in numerous gastropod groups as a variable feature, often associated with gerontic individuals. This does not seem to be the case with the Selsoif species, as most of the specimens are juvenile and both the disjunct whorl and the thicker shell are constant characters.
Wood (1848) described
We are unsure if these Selsoif specimens represent an unknown
-
-
(Fig. 10 H1, H2, I1, I2, J1, J2)
et al. 2011 : 17, pl. 6, fig. 8. — Van Dingenen et al. 2016 : 123, pl. 4, fig. 5. — Van Nieulande et al. 2022 : 20, figs 18-22.
For more, see synonymy list in Fehse & Landau (2003).
Maximum height 12.4 mm, width 9.1 mm. — RGM.1365271 (1), leg. WG; RGM.1365272 (1), leg. WG; RGM.1365003 (42), leg. ACJ; RGM.1365322 (17), leg. AWJ; RGM.1365203 (1), leg. WG; RGM.1365320 (4), leg. WG; RGM.1365274 (8), leg. ACJ; RGM.1365275 (3), leg. AWJ; RGM.1365321 (3), leg. AWJ; RGM.1365057 (20), leg. WG; RGM.1365323 (18), leg. AWJ; RGM.1365105 (239), leg. AWJ.
Aperture well to the right of the ventral mid-line, the fossula not delimited from the columella and the inner border of the columella; fossula not covered by outer lip when seen in ventral view.
Lower Pliocene: NSB, Coralline Crag, England (Wood 1848; Harmer 1920); Atlantic, NW France ( Van Dingenen et al. 2016) ; S Spain ( Landau et al. 2011) . — Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1920), Lillo Formation, Belgium (Glibert 1958; Marquet 1998); Atlantic, Portugal (Silva 2001); western Mediterranean, Spain (Fehse & Landau 2003); central Mediterranean, Italy (Sacco 1894). — Upper Pliocene-Pleistocene: NW France (Brébion 1964). — Lower Pleistocene: Atlantic, Selsoif, France (this paper).
The species varies slightly in the development of the dorsal hump, and in the number of dorsal ribs and labial teeth. This same variability is seen in the Selsoif material, where this species is abundant. We consider this to be the only et al. (2016 : 123) and Van Nieulande et al. (2022 : 20).
-
-
(Fig. 11 A1-A3)
et al. 2022 : 22, figs 33-34.
non J.E Gray, 1832). — Harmer 1920: 512, pl. 45, fig. 2.
et al. 1956 : 85, pl. 10, fig. 100.
Erata (Eratopsis) pernana
Maximum height 5.2 mm, width 3.6 mm. — RGM.1365106 (2), leg. AWJ; RGM.1365287 (1), leg. AWJ.
Shell small, squat, smooth, relatively low spire, outer lip thickened, finely denticulate at inner edge, single terminal fold, two strong anterior columellar denticles, three or four posterior columellar denticles weakening adapically.
Lower Pliocene: NSB, Coralline Crag, England (Wood 1848; Harmer 1920). — Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1920), Belgium (Glibert 1958; Marquet 1997). — Pliocene-Lower Pleistocene (indeterminate): NSB, Netherlands (Beets 1946; Van Regteren Altena et al. 1956 ; Van Nieulande et al. 2022) . — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
The specimens from Selsoif agree with those illustrated by Van Nieulande et al. (2022 : figs 33-34).
-
-
(Fig. 11 B1-B3)
Maximum height 9.2 mm, width 8.3 mm. — RGM.1364921 (188), leg. WG; RGM.1365004 (200+), leg. ACJ; RGM.1365218 (1), leg. AWJ; RGM.1310339 (1), leg. AWJ; RGM.1365107 (500+), leg. AWJ; RGM.1365265 (1 damaged operculum), leg. AWJ.
Globular smooth shell. Three squared teleoconch whorls separated by deep suture. Base covered by a large collumelar lip. Large umbilicus, closed by a thick columellar callus.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
At present, two closely similar extant
Material from the Naturalis collections was re-examined (RP). Five specimens from northern Norway and Nova Zembla gave a first half whorl diameter of 0.44 to 0.48 mm (mean 0.46), agreeing with Bouchet & Warén’s
It is therefore likely that this group is composed of a species complex of more than two species. Detailed morphometric and genetic work is needed to clarify the taxonomic status of the Recent species, which is beyond the scope of this work. We also prefer not to erect a new taxon based on the Selsoif fossil specimens.
-
(Fig. 11 C1-C3)
For more, see synonymy list in Marquet (1997).
Maximum height 26.2 mm (incomplete), width 28.5 mm. — RGM.1365192 (10), leg. WG; RGM.1365193 (6), leg. ACJ; RGM.1365108 (29), leg. AWJ; RGM.1365219 (1), leg. AWJ; RGM.1365273 (2), leg. WG; RGM.1365220 (1), leg. WG; RGM.1365282 (11), leg. ACJ; RGM.1365288 (43), leg. AWJ.
Smooth naticiform shell, with low spire; protoconch consisting of 1.75-2.0 whorls; four convex teleoconch whorls separated by deep suture; last whorl 78% of the total height; aperture large, funicle and inner furrow absent, umbilicus open, with spiral sculpture within.
Lower Pliocene: NSB, Belgium (Marquet 1997; Marquet & Landau 2006). — Upper Pliocene: NSB, Belgium (Marquet 1997), Red Crag, England (Harmer 1921). — Lower Pleistocene: Atlantic, England (Harmer 1921), Sesoif, NW France (this paper). — Upper Pleistocene: NSB, Eemian, Netherlands (Spaink 1958), Atlantic, England, Scotland, Ireland (Harmer 1921). — Holocene: Netherlands (Pouwer & Rijken 2022). Today this species occurs from North Sea (Pouwer & Rijken 2022) to the Mediterranean Sea (Fretter & Graham 1981). We cannot confirm Fretter & Graham’s Mediterranean records and note that Pedriali & Robba (2009) did not record it from the Mediterranean Pliocene.
In their excellent review of the Italian Pliocene Poliniceinae, Pedriali & Robba (2009) characterised
-
-
(Fig. 11 D1-D3)
et al. 2014 : 194, fig. 1. — Símonarson & Eiríksson inEiríksson & Símonarson 2021: 277, pl. 7.5, fig. 2.
For more, see synonymy list in Marquet (1997) and Símonarson et al. (2021) .
Maximum height 32.1 mm, width 18.7 mm. — RGM.1364922 (2), leg. WG; RGM.1365005 (3), leg. ACJ; RGM.1365109 (11), leg. AWJ; RGM.1365187 (1), leg. AWJ.
Shell large, relatively fragile, broadly fusiform; protoconch not preserved; teleoconch of four strongly convex whorls separated by impressed linear suture; sculpture of weak spirals cords with secondaries variably developed; sinuous axial ribs variably developed on early whorls weakening abapically, obsolete on last 1-2 whorls; aperture large, ovate; outer lip weakly thickened by varix, sinuous in profile; columella twisted at fasciole, columella and parietal callus moderately expanded over base, variably thickened.
Lower Pliocene: NSB, Coralline Crag, England (Wood 1848; Harmer 1914). — Upper Pliocene: NSB, Belgium (Marquet 1997), Red Crag, England (Wood 1848; Harmer 1914). — Lower Pleistocene: Atlantic, Iceland (Eiriksson & Símonarson 2021), Atlantic, West Greenland (Símonarson 1981), Selsoif, NW France (this paper). — Upper Pleistocene: western Mediterranean, Spain (Domènech & Martinell 1982); central Mediterranean, Italy (Malatesta & Zarlenga 1986), Baltic ( Funder et al. 2002) . — Holocene: NSB (Strand Petersen 2004). Today, this species is present in the North Atlantic, from Greenland and New Jersey on the western side and Iceland to the Bay of Biscay (Fretter & Graham 1984). Occurs in every kind of habitat, but prefers soft bottoms, lowest tide line to 1200 m depth.
Although highly variable, it cannot be confused with any other species. The Selsoif specimens are mostly incomplete but ascribed to the species without doubt. This is a boreal species.
-
(Fig. 11 E1-E3)
partim): 367, pl. 36, figs 30-31 [not pl. 16, fig. 6, that corresponds to informis Harmer, 1914] (non Harmer, 1914).
partim): 368, pl. 36, fig. 26 [not fig. 28, that corresponds to
et al. 2014 : 31, figs 24-26. — Wesselingh et al. 2014 : 196, fig. 10. — Preece et al. 2020 : fig. 7e.
Maximum height 50.2 mm, width 31.9 mm (incomplete). — RGM.1364923 (18), leg. WG; RGM.1365006 (26), leg. ACJ; RGM.1365188 (1), leg. ACJ; RGM.1310340 (1), leg. ACJ; RGM.1365110 (1), leg. AWJ.
Relatively small, s inistral
Lower Pleistocene: NSB, Norwich Crag and Weybourne Crag, England (Harmer 1918; Preece et al. 2020) ; Atlantic, Selsoif, NW France ( Vervoenen et al. 2014) . — Lower-Middle Pleistocene: Channel region, England ( Vervoenen et al. 2014) . — Middle Pleistocene: Irish Sea (Harmer 1918). — Pleistocene (indeterminate): Netherlands ( Vervoenen et al. 2014) .
et al. 2014) . Those authors noted than the specimen from Selsoif has slightly more convex whorls than usual for specimens of lapsus). Adult specimens are uncommon in Selsoif where the species is represented by hundreds of juveniles. However, one of the specimens illustrated (Pl. 6, fig. 9) has a low spire and very flat whorls in keeping with the species.
-
Based on molecular phylogenetics, all European Neogene nassariids belong in the genus et al. 2016 ).
(Fig. 11 F1-F3)
lapsus, figs 22, 23). non
et al. 2009 : 33, pl. 6, figs 10-12. — Rijken & Pouwer 2014: 49, fig. 19. — Van Dingenen et al. 2015 : 94, pl. 7, fig. 1.
For more, see synonymy list in Landau et al. (2009) and Van Dingenen et al. (2015) .
Maximum height 11.5 mm, width 6.7 mm. — RGM.1364924 (299), leg. WG; RGM.1365209 (1), leg. WG; RGM.1365008 (178), leg. ACJ; RGM.1365296 (1), leg. AWJ; RGM.1008174 (1), leg. AWJ; RGM.1008175 (100+), leg. AWJ.
Shell fusiform, with rounded whorls, separated by deeply impressed, undulating suture. Sculpture of 11-15 rounded, undulating axial ribs, overrun by seven spiral cords per whorl. Outer lip strongly thickened by varix, bearing about seven irregular denticles within. Columella moderately expanded, bearing numerous rugae and tubercles, strong parietal tubercle. Siphonal fasciole broad, separated from base by broad groove.
Lower Pliocene:?central Mediterranean, Italy (Chirli 2000; needs confirmation). — Upper Pliocene: NSB, Red Crag, England (Harmer 1914, 1918); western Mediterranean, S Spain ( Landau et al. 2009) . — Lower Pleistocene: Atlantic, Selsoif, NW France ( Van Dingenen et al. 2015) , Sicily (NHMW coll.). — Middle Pleistocene: Wexford Gravels, Irish Sea (Harmer 1918); Netherlands (Rijken & Pouwer 2014). — Upper Pleistocene: western Mediterranean, Balearic Islands (Cuerda Barceló 1987). Today it is distributed in the eastern Atlantic from Iceland, Northern Norway to the Mediterranean, Canaries (Poppe & Goto 1991), intertidal to 200 m.
For discussion see Landau et al. (2009 : 34) and Van Dingenen et al. (2015 : 94). This is one of the most abundant molluscs in the Selsoif assemblage.
(Fig. 12 A1-A3)
et al. 2009 : 29, pl. 5, figs 15-18, pl. 6, fig. 1, pl. 17, fig. 5. — Chirli & Linse 2011: 160, pl. 54, fig. 3. — Rijken & Pouwer 2014: 54, fig. 35. — Van Dingenen et al. 2015 : pl. 8, fig. 1.
For more, see synonymy list in Landau et al. (2009) and Van Dingenen et al. (2015) .
Maximum height 24.3 mm, width 12.3 mm. — RGM.1364925 (2), leg. WG; RGM.1365204 (1), leg. ACJ; RGM.1365007 (4), leg. ACJ; RGM.1310337 (1), leg. AWJ; RGM.1365111 (40), leg. AWJ.
Shell solid, bucciniform, with elevated conical spire. Protoconch not preserved. Teleoconch of five weakly convex whorls, separated by a shallowly impressed suture. Sculpture of low axial ribs overrun by strap-like cords swollen over the intersection giving surface reticulated appearance. Aperture pyriform; outer lip thickened by varix, denticulated within; columella bearing two folds abapically and parietal fold adapically; columellar callus thickened; parietal callus extended over venter forming parietal shield.
Lower Pliocene: Atlantic, SE Spain (González-Delgado 1989; Landau et al. 2011) ; western Mediterranean, NE Spain (Gili 1991), SE France (Fontannes 1879, Cossmann 1901); central Mediterranean, Italy (Bellardi 1882; Chirli 2000), Tunisia (Fekih 1975). — Upper Pliocene: Atlantic, Portugal (Zbyszewski 1959; Gili et al. 1995 ; Silva 2001); western Mediterranean, S Spain ( Landau et al. 2009) ; central Mediterranean, Italy (Bellardi 1882; Malatesta 1974). — Upper Pliocene-Lower Pleistocene: Atlantic, NW France ( Van Dingenen et al. 2015) . — Lower Pleistocene: Atlantic, Selsoif, NW France ( Van Dingenen et al. 2015) ; central Mediterranean, Italy (Cerulli-Irelli 1911); eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Middle Pleistocene: Atlantic, Wexford Gravels, Irish Sea (Harmer 1916). — Pleistocene (indeterminate): Netherlands (Rijken & Pouwer 2014). — Upper Pleistocene: western Mediterranean, Balearic Islands (Cuerda Barceló 1987). — Holocene: NSB (Strand Petersen 2004). Today, it occurs in the Atlantic and western Mediterranean (Rolán & Luque 1994).
Within the et al. 2009) , et al. (2009 : 30).
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-
(Fig. 12 B1-B3, C1-C3, D1-D2)
et al. 1956 : 88, pl. 11, fig. 114b.
et al 2021: 19, figs 25-29.
Maximum height 54.2 mm, width 33.8 mm. — RGM.1364927 (250+), leg. WG; RGM.1365011 (230+), leg. ACJ; RGM.1365290 (1), leg. ACJ; RGM.1365267 (1), leg. AWJ; RGM.1365114 (500+), leg. AWJ; RGM.1365115 (500+), leg. AWJ; RGM.1405529 (1), leg. R. van Slageren.
Shell solid broadly fusiform; protoconch paucispiral of about 1.5 smooth whorls with large, elevated nucleus. Teleoconch of about three whorls; spire whorls convex, first whorl bearing two primary cords with secondaries intercalated on second whorl; last whorl inflated, with numerous scabrous spiral cords of roughly alternating strength, shoulder and peribasal cords usually slightly stronger; aperture ovate; outer lip not thickened by varix, with internally bevelled edge, with denticles strongly developed to absent; columella thickened abapically, detached, forming medial border of umbilical chink.
Upper Pliocene: NSB, Red Crag, England (Wood 1848; Harmer 1914). — Pliocene and Pleistocene (indeterminate): NSB, Netherlands ( Van Regteren Altena et al. 1956 ; Raad et al 2021). — Lower Pleistocene: NSB, England (Wood 1848; Harmer 1914), Atlantic: Selsoif, NW France (this paper). — Middle Pleistocene: Channel region England (Preece & Bates 1999). — Holocene: NSB (Strand Petersen 2004). Today this species is widely in the northern Atlantic, in the west from Greenland to Connecticut, in the east from Iceland to the Strait of Gibraltar (Fretter & Graham 1984). It inhabits rocky shores to depth of 30-40 m. it lives between 0 and 19°C isotherms (Moore 1936).
et al. 2004 ; inter alia). Some papers have associated these morphs with particular habitats ( Rolán et al. 2004 ) and or presence/absence of predators (Crothers 1983 and others), whereas others have found no association (Berry & Crothers 1974; Crothers 2008). In the Selsoif material, the larger adult specimens usually have spiral cords (Fig. 4 C), smaller but fully adult specimens judging by the thickness of their shell are smoother (Fig. 4 B). The inner lip can be smooth or bear coarse denticles. Some of the juvenile specimens have an imbricated surface with raised lamellae, similar to the juvenile specimen illustrated by Fretter & Graham (1984: fig. 315) as
Similar variability has been described for the size of the protoconch ranging from 800-1400 µm ( Rolán et al. 2004 ) to 500-700 µm (Fretter & Graham 1984). A large number of juveniles are present in the Selsoif material with their protoconch preserved (npw = 1.25-1.5, dp = 740-1000 µm, hp = 730-1030 µm, n= 340-600 µm, p1 = 580-910 µm). These measurements fall between those given above for the recent populations, although the size variation of the nucleus in the Selsoif material is much greater than that reported by Rolán et al. (2004 : table 1). This is a good example of the variability in protoconch size for species with paucispiral protoconchs.
-
(Fig. 12 E1-E3)
cf.
cf.
cf. et al. 2003 26, pl. 7, figs 1-4. — Raad et al. 2021 : 12, figs 1-2.
For more, see synonymy list in Landau et al. (2003) .
Maximum height 9.0 mm, width 4.3 mm. — RGM.1364928 (1), leg. WG.
Small biconic shell. Two smooth protoconch whorls with large bulbous first whorl. Three convex teleoconch whorls separated by an impressed suture. Sculpture on spire whorls of two spiral cords and around ten ribs. Last whorl with six primary cords, shoulder cord appears on second half of last whorl and secondaries intercalated in interspaces; long, narrow siphonal canal.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). The oldest stratigraphic record of et al. 2007 , 2011) and western (Martinell & Marquina 1981) and central Mediterranean (Chirli 2000). — In the Upper Pliocene it is present in the Mondego Basin of Portugal (Silva 2001) and is widespread in the Mediterranean. Today it occurs from the southern British Isles to the Azores, Madeira, and Canary Islands and entire Mediterranean (Houart 2001).
We attribute the specimen with some hesitation to
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-
(Fig. 12 F1-F3)
Maximum height 3.6 mm, width 1.4 mm. — RGM.1365116 (6), leg. AWJ; RGM.1365293 (1), leg. ACJ; RGM.1365292 (5), leg. ACJ.
Shell small, slender fusiform. Protoconch not preserved. Teleoconch of 4.5 convex, weakly shouldered whorls bearing ten elevated, opisthocline ribs, about half width of their interspaces. No sign of spiral sculpture; possibly abraded. Last whorl elongate, weakly shouldered, moderately constricted at base. Aperture elongate, outer lip incomplete. Columella weakly excavated, siphonal canal moderately long, open, wide.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
All specimens are too incomplete and abraded to offer further identification.
-
(Fig. 12 G1-G3)
Maximum height 10.7 mm, width 4.4 mm. — RGM.1365012 (90+), leg. ACJ; RGM.1365294 (21), leg. ACJ; RGM.1364929 (31), leg. WG; RGM.1365291 (1), leg. AWJ; RGM.1365117 (115), leg. AWJ.
Small fusiform shell. Two smooth protoconch whorls. Three quadrate teleoconch whorls separated by well-defined swallow suture. Sculpture consists of two spiral cords conferring the quadrate aspect of the whorls; and around 15 opisthocline axial ribs. Last whorl is covered by numerous spiral cords and ribs, continuing on the base. Well developed columelar lip. Elliptical aperture, with a short, widely open siphonal canal.
Lower Pleistocene: Atlantic, Selsoif (this paper).
Even though et al. 2016) , the elongate-shaped specimens with well pronounced ribbing from Selsoif appear to fall outside the range of variation. Further study is required to establish whether this might represent a new species.
-
-
(Fig. 12 H1-H3)
Murex linearisMontagu, 180 3: 261, pl. 9, fig. 4.
sic ])
et al. 2022 :170, pl. 10, figs 1-3.
For more, see synonymy list in Landau et al. (2022) .
Maximum height 9.4 mm, width 3.6 mm. — RGM.1364930 (17), leg. WG; RGM.1365013 (43), leg. ACJ; RGM.1365215 (1), leg. ACJ; RGM.1365118 (50), leg. AWJ; RGM.1365215 (1), leg. ACJ.
Small, fusiform shell. Protoconch not preserved. Five convex teleoconch whorls separated by weakly marked suture. Sculpture consists of 12 orthocline to prosocline, broad, round ribs and two spiral cords running over the ribs on the first teleoconch whorl, the third appears below the suture and the fourth above the suture on second whorl. Third whorl has 5 spiral cords. Last whorl with six spiral cords, continuing until the columella. Aperture ovate, extended by a broad, twisted siphonal canal.
Upper Miocene: Atlantic, southwestern Spain ( Cárdenas et al. 2019) ; central Proto-Mediterranean, Italy (Montanaro 1937: Venzo & Pelosio 1963). — Lower Pliocene: NSB, Coralline Crag, England (S.V. Wood 1848; Harmer 1915); Atlantic, Guadalquivir Basin, S. Spain (González-Delgado 1993); central Mediterranean, Italy (Chirli 1997; Brunetti & Cresti 2018). — Upper Pliocene: NSB, Red Crag, England (S.V. Wood 1848; Harmer 1915); western Mediterranean, Estepona Basin, southern Spain (Vera-Peláez 2002; Landau et al. 2022) ; central Mediterranean (Bellardi 1877; Cavallo & Repetto 1992). — Lower Pleistocene: central Mediterranean, Italy (Cerulli-Irelli 1914). — Middle Pleistocene: eastern Mediterranean, Rhodes Island (Chirli & Linse 2011). — Pleistocene (indeterminate): Atlantic, England (Harmer 1915, 1918). — Upper Pleistocene: western Mediterranean, Balearic Islands (Cuerda Barceló 1987). Holocene: NSB (Strand Petersen 2004). — Present-day: Atlantic: Norway (Høisæter 2016), Scotland (E.H. Smith 1967), British Isles (Fretter & Graham 1984; Graham 1988), Galicia ( Oliver et al. 2022) , south to Portugal (Nobre 1931) and Canary Islands (Nordsieck & García-Talavera 1979); western Mediterranean ( Bucquoy et al. 1882 ), central Mediterranean ( Bogi et al. 1980 ; Cachia et al. 2001) . Today it is occurs in the Atlantic from Norway (Høisæter 2016), Scotland (Smith 1967) and British Isles (Fretter & Graham 1984; Graham 1988) to Galicia ( Oliver et al. 2022) , south to Portugal (Nobre 1931) and Canary Islands (Nordsieck & García-Talavera 1979) and in the western Mediterranean ( Bucquoy et al. 1882 ) and central Mediterranean ( Bogi et al. 1980 ; Cachia et al. 2001) .
For discussion see Landau et al. (2022) .
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-
(Fig. 13 A1-A3, B1-B3)
urn:lsid:zoobank.org:act:FD2A03A9-3782-423B-9D89-E5BC08BDE470
MNHN.F.A91318, height 5.4 mm, width 2.9 mm, leg. SM.
Maximum height 8.7, width 5.2. — RGM.1365014 (1), leg. ACJ.
Name reflecting the close resemblance to members of the cancellariid genus
Selsoif, Saint-Sauveur-le-Vicomte, Manche department, NW France.
Gelasian, Early Pleistocene
Medium sized
The Selsoif specimens are relatively consistent in profile and sculpture, although in some the subsutural ramp is slightly slanted giving the profile a more typically Admete-like appearance. The width and concavity of the subsutural ramp is somewhat variable. In one specimen, a further cord runs on the subsutural platform just medial to the shoulder cord on the last two whorls and a fine secondary spiral is intercalated between the second and third primary cords. The subobsolete axial sculpture in this same specimen is slightly stronger, forming poorly defined axial ribs. The small size of the specimens and the unthickened outer lip suggests that they may not represent fully grown adults.
Lower Pleistocene: Atlantic, endemic to Selsoif (this paper).
Shell small and relatively thin for genus, with moderate height scalate spire. Protoconch paucispiral, dome-shaped, about 1.5 whorls, surface abraded. Teleoconch of four low, barrel-shaped, strongly tabulate whorls, separated by narrowly impressed, linear suture: subsutural platform broad, increasingly concave and canaliculate abapically, delimited by elevated shoulder cord. Sculpture on early teleconch whorls of three narrow spiral cords, with fourth appearing just above suture. On penultimate whorl single secondary cord intercalated between shoulder cord and mid cord. Axial sculpture subobsolete, represented mainly by indistinct tubercles over cords, and crowded slightly prosocline growth-lines. Last whorl 70% of total height with relatively broad, shallowly canaliculate subsutural platform, sharply angled at shoulder cord, weakly constricted at base, sculptured of four weakly tubercular cords above level of insertion of outer lip, about eight further smooth cords below, becoming closer-spaced over siphonal fasciole; base not delimited; siphonal fasciole weakly developed, slightly rounded with very narrow umbilical chink medially. Aperture 50% of total height, subquadrangular, outer lip not thickened, sharply angled at shoulder; anal sinus lacking, siphonal canal moderately short, open, bent adaxially. Columella excavated in upper third, straight below, twisted at fasciole, bearing three columellar folds of subequal strength, becoming increasingly oblique abapically. Columellar callus thickened, sharply delimited medially by umbilical chink, parietal callus not developed.
The strongly scalate spire and deeply canaliculate suture are reminiscent of members of the cancellariid genus et al. (2011 : table 1) as a thermophilic marker, which has its maximum diversity in tropical assemblages and does not occur in waters that are colder than subtropical.
Placement in the temperate to cold water genus
-
(Fig. 13 C)
For more, see synonymy list in Landau & Micali (2021).
Maximum height 0.5 mm, width 0.2 mm. — RGM.1365080 (1), leg. AWJ.
Small, thin turriform shell. Protoconch and early teleoconch whorls not preserved. Three convex teleoconch whorls, swollen in abapical half. Sculpture of sinuous axial ribs and fine spiral cords in visible in axial interspaces.
Lower Pliocene: central Mediterranean, Italy (Chirli & Bogi 2002; Chirli 2013). — Upper Pliocene: Atlantic, Mondego Basin, Portugal (Landau & Micali 2021); western Mediterranean, Estepona Basin, S. Spain (Landau & Micali 2021). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). Today this species occurs in the eastern Atlantic of south Norway, British Isles (Fretter & Graham 1982), North Sea (Raven 2020) to Canaries ( Hernández et al. 2011) , Madeira and Selvagens Islands ( Segers et al. 2009) , Cape Verde Islands, West Africa (Rolán 2005) and Mediterranean, Italy ( Giannuzzi-Savelli et al. 2014) .
-
-
(Fig. 13 D1-D3)
et al. 1965 : 26, pl. 21, fig. 205. — Ceulemans et al. 2018 : 122, pl. 7, fig. 9.
For more, see synonymy list in Ceulemans et al. (2018) .
Maximum height 1.7 mm, width 0.8 mm. — RGM.1365120 (20), leg. AWJ; RGM.1365121 (1), leg. AWJ; RGM.1365017 (3), leg. ACJ.
Shell small, subcylindrical with low spire. Two to three teleoconch whorls separated by deeply impressed suture. Surface smooth. Aperture elongate expanded abapically.
Lower Pliocene: NSB, England (Harmer 1923); Atlantic, NW France ( Ceulemans et al. 2018) . — Upper Pliocene: NSB, Red Crag, England (Wood 1848). — Upper Pliocene-Pleistocene: NSB, Holland ( Van Regetren Altena et al. 1955 ). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper). — Middle Pleistocene: Atlantic, Irish Sea (Harmer 1923). Today, this species occurs in the Atlantic from Greenland, Iceland to Scandinavia, Nova Scotia to Aleutian Islands, British Isles (Thompson 1988).
et al. 2018) . As noted by those authors, this species is rarely reported in the fossil literature. It is predominantly a cooler water species with the most southern record being that of the Lower Pliocene of Le Pigeon Blanc (NW France).
-
-
(Fig. 13 E)
Fragment of apex. — RGM.1365018 (1), leg. ACJ., W of incomplete fragment 3.5 mm.
Apical fragment.
This single fragment probably represents a
-
-
(Fig. 13 F1-F3)
cf.
cf.
For more, see synonymy list in Landau & Micali (2021).
Maximum height 2.9 mm (incomplete), width 1.3 mm. — RGM.1365276 (1), leg. AWJ.
Three smooth, weakly convex abraded adult whorls preserved. Aperture broken. Columella marked by three folds; abapical two folds weaker.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
This fragment is unusual in having columellar folds suggesting placement in the genus
-
(Fig. 13 G1-G3)
For more, see synonymy list in Landau & Micali (2021).
Maximum height 1.1 mm, width 0.6 mm. — RGM.1365020 (2), leg. ACJ; RGM.1365214 (1), leg. AWJ; RGM.1365122 (50+), leg. AWJ.
Small shell with tall conical spire, type A2 helicoid protoconch (Landau et al., 2020a), almost flat-sided teleoconch whorls, last whorl angular at the periphery, prosocline growth-lines and well developed columellar fold.
Middle Miocene: Proto-Mediterranean Sea (Serravallian): Karaman Basin, Turkey ( Landau et al. 2013) . — Upper Miocene: Atlantic (Tortonian and Messinian), NW France ( Landau et al. 2020a) . — Lower Pliocene: NSB, Coralline Crag, England (Harmer 1923); western Mediterranean, Tunisia (Fekih 1969); central Mediterranean, Italy (Sacco 1892; Crovato & Micali 1992b; Guioli et al. 2009 ; Chirli & Micali 2011). — Upper Pliocene: Atlantic, St Erth, England (Harmer 1923), NSB, Red Crag, England (Harmer 1923), Kruisschans and Oorderen Sands, Belgium (Marquet 1993, 1998); Atlantic, Mondego Basin, Portugal (Landau & Micali 2021); western Mediterranean, Estepona Basin, S. Spain (Peñas & Rolán 1999; Landau & Micali 2021); central Mediterranean, Italy (Sacco 1892;? Cavallo & Repetto 1992; Ferrero et al. 1998) . — Upper Pliocene-Pleistocene: NSB, Netherlands ( Van Regteren Altena et al. 1964 ). — Lower Pleistocene: Selsoif, NW France (this paper); central Mediterranean, Italy (Cerulli-Irelli 1914; Gianolla et al. 2010 ; Brunetti 2011); eastern Mediterranean, Rhodes (Chirli & Linse 2011). — Upper Pleistocene: Ireland (Harmer 1923). — Present-day: eastern Atlantic from North Iceland, Norway to British Isles (Nordsieck 1972; Høisæter 2014), North Sea (Raven 2020), northwestern Spain ( Trigo et al. 2018) , Azores, Madeira and Selvagens Islands ( Segers et al. 2009) , south to Canaries ( Hernández et al. 2011) , West Africa, Cape Verde Islands to Angola ( Peñas et al. 2009 , 2014), into western Mediterranean ( Bucquoy et al. 1882 ; Peñas et al. 1996) , central Mediterranean ( Cachia et al. 2001 ; Giannuzzi-Savelli et al. 2014) , eastern Mediterranean ( Öztürk et al. 2013) .
(Fig. 13 H1-H3)
Maximum height 2.2 mm, width 1.4 mm. — RGM.1365098 (10), leg. AWJ; RGM.1365199 (1), leg. AWJ; RGM.794508 (c. 250), leg. AWJ; RGM.961907 (1), leg. AWJ.
Thick, smooth, ovate shell. Protoconch heterostrophic. Two teleoconch whorls separated by a deep suture. Pyriform aperture with small fold deep within columella.
This species is unusual in being extremely small and thick-shelled. The protoconch is clearly heterostrophic and there is a small columellar fold deep within the aperture seen only in tangential view. We are unsure where to place this species and leave it in open nomenclature.
Lower Pleistocene: Atlantic, Selsoif (this paper).
-
(Fig. 13 I1-I3)
Maximum height 1.1 mm, width 0.6 mm. — RGM.1365021 (1), leg. ACJ; RGM.1365266 (1), leg. AWJ; RGM.794507 (19), leg. AWJ; RGM.961906 (1), leg. AWJ; RGM.1365022 (1), leg.ACJ, RGM.1365123 (6), leg. AWJ.
Elongate-ovoid shell, strongly intorted type C protoconch ( Landau et al. 2020a) , teleoconch of three moderately convex whorls, separated by deeply impressed suture, opisthocline to orthocline growth-lines, sinuous towards adapical suture, fine spirals cover entire whorl, last whorl inflated, regularly convex, narrow umbilical chink. Aperture pyriform, columella bearing weak fold mid-aperture.
Lower Pliocene: NSB, Coralline Crag, England (Wood 1872); Kattendijk Formation, Belgium (Marquet 1997). — Late Pliocene: St Erth, Atlantic, England (Harmer 1923), Selsoif, NW France (this paper). Today this species occurs in northern Europe from North Norway to the Bay of Biscay ( Fretter et al. 1986 ).
The specimens from Selsoif differ slightly from modern ones in having the spiral sculpture extending along the entire whorl surface, whereas in extant specimens it is restricted to the abapical portion of the whorl.
(Fig. 14 A1-A3)
cf.
cf.
et al. 2020c : 298, pl. 22, figs 1-3. — Landau & Micali 2021: 298, pl. 22, figs 1-3.
Maximum height 2.1 mm, width 0.9 mm. — RGM.1365019 (4), leg. ACJ; RGM.1365213 (1), leg. ACJ.
Small, solid shell. Protoconch not preserved. Five teleoconch whorls, spire telescopic, whorls weakly convex, surface appears smooth (abraded). Outer lip prosocline in profile and arched. Small columellar fold. Narrow umbilical chink.
Upper Miocene: Atlantic, NW France ( Landau et al. 2020a) . — Upper Pliocene: western Mediterranean, Estepona Basin, Spain (Landau & Micali 2021). — Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
This seems to be the same species as that recorded by Landau et al. (2020a) from the Upper Miocene of NW France and Landau & Micali (2021) from the Upper Pliocene Estepona Basin of Spain as
-
(Fig. 14 B1-B3)
Maximum height 1.6 mm, width 0.8 mm. — RGM.1365207 (1), leg. AWJ.
Shell fusiform. Protoconch type C. Teleoconch of 3.5 convex whorls separated by narrowly impressed suture. Sculpture consist of numerous spiral cords, broader than their interspaces, six on first teleoconch whorl, 18 on last whorl. Last whorl 70% of the total height. Aperture pyriform; outer lip not complete; columella without fold.
Lower Pleistocene: NSB, England (Wood 1879; Harmer 1920); Atlantic, Selsoif, NW France (this paper).
We tentatively ascribe the specimen to
-
(Fig. 14 C1-C3)
For more, see synonymy list in Landau & Micali (2021).
Maximum height 1.2 mm, width 0.6 mm. — RGM.1365024 (5), leg. ACJ; RGM.1365126 (450+), leg. AWJ; RGM.1365205 (1), leg. AWJ.
Small turriform shell. Flat protoconch of one whorl. Teleoconch of 5 flat-sided whorls separated by canaliculated suture with 20 orthocline, axial ribs stopping at adapical cord. Base marked by weakened spiral cord. High, pyriform aperture with one tooth on the inner lip, at the level of a small umbilicus.
First occurences in the Upper Miocene from the Atlantic, NW France ( Landau et al. 2020a) . In the Lower Pliocene, only present in the Mediterranean of Italy (Sacco 1892; Ferrero Mortara et al. 1984 ; Crovato & Micali 1992a; Bogi & Chirli 2004; Chirli & Micali 2011). This species occurs in the Upper Pliocene from the Mediterranean of Estepona Basin, Spain (Landau & Micali 2021), Italy (Crovato & Micali 1992b; Ragaini & Bernieri 2007). Present in the Lower Pleistocene from the Mediterranean of Italy (Crovato & Micali 1992a) and Rhodes (Chirli & Linse 2011). Last fossil occurence in the Upper Pleistocene from Mediterranean, Italy (Crovato & Micali 1992a). Today this species occurs in the Atlantic from northwestern Spain ( Trigo et al. 2018) , south Portugal, Canaries ( Hernández et al. 2011) to West Africa (Peñas & Rolán 1998; Van Aartsen et al. 2000 ; Peñas et al. 2014) , and in the Mediterranean ( Peñas et al. 1996 ; Giannuzzi-Savelli et al. 2014 ; Micali & Palazzi 1992; Öztürk et al. 2011) to the Black Sea (Nordsieck 1972; Wilke & Van Aartsen 1998).
-
(Fig. 14 D1-D3)
Maximum height 4.3 mm (incomplete), width 1.5 mm. — RGM.1364932 (1), leg. WG; RGM.1365023 (68), leg. ACJ; RGM.794509 (c. 95), leg. AWJ.
Fragment consisting of last 4.5 whorls, sculptured by broad, curved, opisthocline ribs, wider than their interspaces, stopping abruptly at suprasutural cord and at base.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
The character of the spiral sculpture stopping abruptly at a suprasutural cord on the later spire whorls and at the base on the last whorl allows attribution to the genus
-
(Fig. 14 E1-E3)
Maximum height 1.3 mm, width 0.6 mm. — RGM.1365304 (1) leg. AWJ; RGM.1365305 (1), leg. AWJ; RGM.1365223 (4), leg. AWJ.
Small, relatively broad turriform shell. Protoconch absent. Last 4.5 teleoconch whorls preserved, whorls rather flat-sided, first two whorls surface abraded, last two whorls with very broad opisthocline ribs separated by narrow interspaces. Pyriform aperture angled at top of straight columella. Subobsolete columellar fold.
Lower Pleistocene: Atlantic, Selsoif, NW France (this paper).
The species from Selsoif has a similar outline to that of
- indet.
(Fig. 14 F1-F3)
Height 1 mm, width 0,52 mm. — RGM.1365029 (1), leg. AWJ; RGM.1365324 (1), leg AWJ, RGM.1310835 (1), leg. AWJ; RGM.1364976 (1), leg. AWJ.
Small turriform shell. Protoconch heterostrophic. Three convex teleoconch whorls separated by a weakly impressed suture. Sculpture of two spiral cords on spire whorls, with numerous lamellar axial riblets in spiral interspaces. Last whorl with three spiral cords, abapical cord delimiting base, cords broaden and weaken abapically, base rounded, riblets weaken on last whorl, subobsolete. Aperture pyriform, no columellar fold.
Lower Pleistocene: Atlantic, endemic to Selsoif (this paper).
This single specimen has very distinctive sculpture of strong spirals and riblets in the interspaces. It might represent an
-
- indet.
(Fig. 14 G1-G3)
et al. 2018 : 140, pl. 9, figs 3-5.
For more, see synonymy list in Ceulemans et al. (2018) .
Maximum height 23.5 mm, width 13.2 mm. — RGM.1364933 (7), leg. WG; RGM.1365025 (11), leg. ACJ; RGM.1365127 (15), leg. AWJ; RGM.1365208 (1), leg. AWJ.
Shell medium sized, globular, teleoconch composed of about eight smooth whorls, except for subsutural cord (abraded in most specimens); spire whorls flat-sided, last whorl inflated, aperture elongate-ovate with two narrow columellar teeth, narrow, deep umbilicus.
Lower Pliocene: NSB, Red Crag, England (Harmer 1923); Atlantic, NW France ( Ceulemans et al. 2018) . — Upper Pliocene: Atlantic, St Erth, England (Harmer 1923); NSB, Red Crag, England (Wood 1848; Harmer 1923), Oorderen Sands, Belgium (Marquet 1997). — Pliocene (indeterminate): NSB, Netherlands ( Van Regteren Altena et al. 1964 ). — Upper Pliocene-Pleistocene: Atlantic, NW France (Brébion 1964). — Lower Pleistocene: Selsoif, NW France (this paper).
This is the only terrestrial pulmonate gastropod in the Selsoif fauna. Ellobiids typically live in supratidal habitats. This species has usually been placed in the genus et al. (2023) showed that
The gastropod fauna of Selsoif contains 76 species-level taxa (Table 1). Abundant species are viz.
The studied fauna contains a number of groups with very different biogeographic affinities (Table 1). The vast majority of the fauna consists of extant species, all of which live in the Boreoceltic, Franco-Iberian to Mediterranean realm. This component has a distinct modern littoral character and includes species such as
The Selsoif fauna is predominantly a modern East Atlantic littoral fauna, but with some transitional components. The fauna has almost no species in common with the Tortonian-Zanclean faunas from the lower Loire and Brittany region of Assemblages I-III (of Van Dingenen et al. 2015 ; commonly referred to as the Redonian). Their Assemblage IV (Upper Pliocene-Pleistocene) may share more species in common with that of Selsoif but is at present too poorly known to be certain (BL personal observation). Although the Selsoif assemblage has previously been associated with the ‘Redonian’ (e.g., Marquet 2001), the latter term is now restricted to Miocene-Pliocene associations in the Ligerian Gulf (see Landau et al. 2020b) and should not be applied to Selsoif.
There are some species in common with the latest Pliocene St Erth Beds from Cornwall and the Bosq d’Aubigny fauna from the Cotentin, but both faunas represent different, calmer depositional settings. Furthermore, both the St Erth and the Bosq d’Aubigny faunas require revision and further description to enable meaningful comparisons with the Selsoif fauna. The Selsoif fauna has several species in common with Late Pliocene and especially Early Pleistocene faunas from the North Sea Basin. Species in common with the Early Pleistocene Norwich and Weybourne Crag Formations of eastern England and the Dutch Maassluis Formation are et al. 2007 ; Preece et al. 2020) .
Remarkably, three extinct species are present in the Selsoif fauna that were known previously only from North Sea Basin Pliocene, viz. et al. 2021 ; Wesselingh et al. 2023) and the south coast of England. (Harmer 1917). It concerns common species such as
The Gelasian gastropod fauna from Selsoif (Cotentin, France) represents a diverse marine littoral fauna that is dominated by extant species but also contains a few endemics and some Pliocene relict species. The biogeographical affinity of the fauna is one that is typical of the southern part of the Boreo-Celtic and northern part of the French-Iberian province but the overall assemblage differs from Late Quaternary/modern assemblages in the region in terms of species composition. The Selsoif fauna therefore forms a transition between Pliocene and modern faunas in the temperate East Atlantic domain and shows that the modern nature of molluscan faunas already existed in the Early Pleistocene.
We are greatful to Han Raven and Riaan Rijken for advice on specific taxonomic groups. We are deeply indebted to Len Vaessen and Stef Mermuijs for making available their material for this project. We thank Didier Merle (MNHN, Paris) for his support of the project. Richard Preece (University Museum of Zoology, Cambridge, United Kingdom) and Mathias Harzhauser (Natural History Museum Vienn, Austria) made the review of the original manuscript and provided very useful comments that helped to improve this paper.
et al.2000 and Pareyn 1987) showing the location of Selsoif:
1, Precambrian-Palaeozoic basement;
2, Permo-Mesozoic Paris Basin fill;
3, Neogene-Quaternary subbasins.
A, photographie IGN, Mission C90SAA0491_1990_F1211- 1411_0037, cliché no. 37, 28/04/1990;
B, photographie IGN, Mission CP12000422_059_12385, cliché n°12385, 26/07/2012. Width of lake in the centre
c.400 m.
et al.2011) .
A,
B,
C,
D,
E,
F,
G,
H,
I,
J,
K,
A,
paratype 1), RGM.1365210, leg. WG, L = 5.2 mm;
B,
C,
holotype), RGM.1362824, H = 580 μm;
D,
E,
F,
G,
H,
I,
J,
A,
B,
C,
holotype), MNHN.F.A91316, leg AWJ, H = 4.2 mm;
D,
paratype), RGM.1365312, leg AWJ, H = 3.9 mm;
E,
F,
G,
A,
B,
C,
D,
E,
F,
G,
H,
I,
J,
A,
B,
holotype), RGM.1365286, leg ACJ, H = 1.28 mm;
C,
D,
E,
F,
A,
B,
C,
D,
E,
F,
A,
B,
C,
D,
E,
F,
G,
H,
I,
J,
A,
B,
C,
D,
E,
F,
A,
B,
C,
D,
E,
F,
G,
H,
A,
B,
C,
D,
E,
F,
Longchaeus
G,
H,
I,
A,
B,
C,
D,
E,
F,
G, “
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