Timbellus occidentalis, Merle & Pacaud & Ledon & Goret, 2024

Merle, Didier, Pacaud, Jean-Michel, Ledon, Daniel & Goret, Bernard, 2024, New Cenozoic Muricidae (Mollusca: Gastropoda) from Europe, Geodiversitas 46 (15), pp. 495-551 : 512-513

Timbellus occidentalis n. sp.

( Figs 8 A-C; 26A, B)

urn:lsid:zoobank.org:act:53A32E50-A05D-4A09-B8B1-6CA776FDC29F

Murex (Pterymurex) tripteroides – Cossmann & Pissarro 1901: 78, pl. 13, fig. 29 [non Lamarck, 1822].

Timbellus cf. tripteroides – Merle et al. 2011: 446, pl. 101, figs 1, 2.

TYPE MATERIAL. — Holotype. France • Normandy, Manche, Fresville; Calcaire de Fresville ; middle Eocene (uppermost Lutetian ); MNHN.F.A90533 (ex A24100 ex galerie de zoologie coll., Le Marchand coll.), H: 20 mm ( Figs 8A, B ; 26A ).

Paratypes. France • 1 spm; same as for the holotype; MNHN.F.A90534 (ex A24100 ex galerie de zoologie coll., Le Marchand coll.) • 4 spm; idem; MNHN.F.B63570 (de Morgan coll.) • 1 spm, MNHN.F.A90532 (de Morgan coll.) • 1 spm; Normandy, Manche, Hautteville-Bocage; middle Eocene (upper Lutetian); MNHN.F.A90531 (ex B63609, de Morgan coll.) • 1 spm; idem; MNHN.F.B63609 (de Morgan coll.) • 35 spm; idem; MNHN.F.B63610 (de Morgan coll.) – Gourbesville, Manche, middle Eocene (upper Lutetian) • 3 spm; idem; MNHN.F.A24128 (Munier-Chalmas coll.) • 10 spm; Loire Basin, Loire-Atlantique, Saffré (Bois-Gouët); middle Eocene (lower Bartonian); MNHN.F.A29658 (Pacaud coll.) • 1 spm; idem; MNHN.F.A24031 (Vasseur coll.), figured in Merle et al. (2011: pl. 101, fig 1) as Timbellus cf. tripteroides ( Lamarck, 1822) ( Figs 8C ; 26B ) • 1 spm; idem; MNHN.F.J06383 (Vasseur coll.), figured in Merle et al. (2011: pl. 101, fig 2) as Timbellus cf. tripteroides ( Lamarck, 1822) .

ETYMOLOGY. — From the Latin occidentalis meaning west, considering the geographic range of this species in Normandy and in the Loire Basin.

TYPE HORIZON. — Calcaire de Fresville, middle Eocene (uppermost Lutetian), see Bignot et al. 1968).

TYPE LOCALITY. — France, Normandy, Manche, Fresville, see APGN (2024).

DISTRIBUTION. — Normandy (Calcaire de Fresville and Faluns de Hautteville) middle Eocene (uppermost Lutetian) and Loire Basin (Sables du Bois-Gouët), lowermost Bartonian see Mathelin & Bignot (1989).

DESCRIPTION

Smooth, bulbous protoconch of 1.25 whorls. Teleoconch up to 26.6 mm in height, up to 15.3 mm in width, biconic in profile, composed of five whorls. Moderately high spire. Last whorl up to 85% total length. Apical angle 64° excluding spines, 91° including spines. Spiral sculpture with weak primary cords in adult specimens. First to fourth whorl, no cord. Fifth whorl: P1 to P3. Sixth whorl: P1 to P6 weak, s2 and no other cord below, ADP and MP occasional. On first whorl, eight protovarices. From second to last whorl, three varices and one intervarix intercalated Higher relief of intervarices on P2-P3. Varices lamellose, winged on last whorls. Appearance of short, straight P1 spine on third whorl. Sinus of P1 spine narrow and pointing adapically. On last whorl alate varices. Ovate aperture up to 9% of diameter and up to 80% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, slightly erect anteriorly, forming curved inductura at base. Parietal lip slightly adherent. Outer lip with denticles ID (occasional), D1 to D6, D1 and D2 more strongly developed. Outer lip not crenulate. Siphonal canal narrow, open, up to 49% of apertural length. Pseudo-umbilicus narrow.

4

3

cm

in

Width 2

1

0 0 1 2 3 4 5 6 7

Total length in cm

COMPARISONS

This species was previously attributed to Murex tripteroides Lamarck, 1822 by Cossmann & Pissarro (1901) and later with some doubts by Merle et al. (2011). According to those authors, there are apparently strong similarities with T. tripteroides such as: a bulbous protoconch, an elongate shape, developed internal denticles, and few and weak cords. However, T. occidentalis n. sp. is smaller and does not exceed 26 mm for five teleoconch whorls. At the same size and same number of whorls T. tripteroides is still juvenile, does not bear internal denticles, and their axial wings are only developed on the last whorl ( Fig. 8D, E ). Adults of T. tripteroides can reach more than 60 mm height at 7-8 whorls. A bivariate diagram ( Fig. 9 ) illustrates the differences in size between T. occidentalis n. sp. from Normandy and T. tripteroides from the Paris Basin. In addition, the spiral cords of T. tripteroides ( Figs 6 ; 8D, E ) are usually more marked than those of T. occidentalis n. sp. ( Fig. 8 A-C) Finally, T. occidentalis n. sp. could be regarded as a progenetic species of Timbellus , because it associates a small size with adult characters comparable to T. tripteroides . Moreover, the two species never co-occur. Timbellus occidentalis n. sp. ranges between the uppermost Lutetian (Normandy) and the lowermost the Bartonian (Loire-Atlantique), whereas T. tripteroides ranges from the lower to the middle Lutetian in the Paris Basin and in the Hampshire Basin during the Bartonian.

Figures

FIG. 8. — Spiral cords and internal denticles of the outer lip in Timbellus occidentalisn. sp.(A-C) and T. tripteroides (Lamarck,1822) for comparison (D-E): A, B, holotype,MNHN.F.A90533 (Le Marchand coll.), Fresville,Manche,Normandy,France, middle Eocene (upper Lutetian);C, paratype,MNHN.F.A24031 (Vasseur coll.), Saffré (Bois-Gouët), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); D, E, MNHN.F.A90540 (Merle coll.), Beynes (Ferme de l’Orme), Yvelines, Paris Basin, France, middle Eocene (Lutetian). Scale bars: 5 mm. Credits: A-C, P. Loubry; D, E, L. Cazes.

FIG. 26. — New species of Timbellus De Gregorio, 1885: A, B, Timbellus occidentalis n. sp.: A, holotype, MNHN.F.A90533 (Le Marchand coll.), Fresville, France, upper Lutetian;B, paratype,MNHN.F.A24031 (Vasseur coll.),Saffré (Bois-Gouët),France,Bartonian;C, D, T.calciacus n. sp.: C, holotype,MNHN.F.A90536 (Merle coll.), Chaussy (Les Garennes), France, Lutetian; D, paratype, MNHN.F.A90537 (Merle coll.), Chaussy (Les Garennes); E, F, T. longicanalis n. sp.: E, holotype, MNHN.F.A24101 (Le Marchand coll.), Fresville, France, upper Lutetian; F, paratype, MNHN.F.A24100, Fresville; G-I, T. magnei n. name: G, MNHN.F.A32508 (Merle coll.), juvenile specimen, Gan (Tuilerie), France; H, MNHN.F.A90545 (Merle coll.), juvenile specimen with protoconch, Gan (Tuilerie); I, MNHN.F.A72443 (Varone coll.), Gan (Tuilerie); J-L, T. curvispina n. sp.: J, holotype, MNHN.F.A90541 (MNHN coll.), Parnes, France, Lutetian; K, paratype, MNHN.F.A90543 (Pons coll.), Fontenay-en-Vexin, France, Lutetian; L, paratype, MNHN.F.A73848 (Pons coll.), Fontenay-en-Vexin; M, T. sixi n. sp., holotype, MNHN.F.A90544 (Six coll.), Trosly-Breuil (Fond Couturier), France, Ypresian; N-P, T. radulfiensis n. sp.: N, holotype, MNHN.F.A77754 (MNHN coll.), Rauville-la-Place, France, Priabonian; O, paratype, MNHN.F.A77755 (MNHN coll.), Rauville-la-Place; P, paratype, MNHN.F.A90559 (Vasseur coll.), Rauville-la-Place. Scale bars: A,B, N-P, 5 mm; C, D, 3 mm; E-G, I-L, 10 mm; H, 2 mm.

FIG. 9. — Bivariate plot of total length against width comparing Timbellus tripteroides (Lamarck,1822) from the middle Lutetian of Paris Basin (65 complete specimens) and T. occidentalis n. sp. from the upper Lutetian of Normandy (18 complete specimens).

FIG. 6. — Neotype, historical figures, spiral cords and denticles of the outer lip in T. tripteroides (Lamarck, 1822): A-D, morphotype with rounded varices and poorly distinguishable P1 spine; E-H, morphotype with easily distinguishable P1 spine: A, original figure of Lamarck (1805: pl. 3, fig. 4); B, C, neotype designated herein, MNHN.F.A90546 (Club géologique d'Île-de-France leg.), Thiverval-Grignon (Falunière), Yvelines, Paris Basin, France, middle Eocene (Lutetian); D, MNHN.F.A90552 (Pacaud coll.), Chaussy (Les Garennes),Paris Basin, France,middle Eocene (Lutetian); E, F, figured specimen of Deshayes (1835: pl. 82, fig. 1-2); G, H, figured specimen of Cossmann & Pissarro (1911: pl. 35, fig. 169-1; MNHN.F.J10350 (Cossmann coll.), Chaussy (Les Garennes). Scale bars: 20 mm. Credits: B-D, P. Loubry MNHN/CNRS; G, H, J. Mouchart.