Chicoreus(Phyllonotus)initialis ‌Vokes, 1990. United States, Texas, middle Eocene (Bartonian) (Fig. 2A, B).

Murex ‌packardi ‌Dickerson, 1915, Washington (United States), middle Eocene; Chicoreus(Phyllonotus)initialis ‌Vokes, 1990, Texas (United States), middle Eocene (Lutetian); Pterynoyus newtoni Eames, 1957 [new name for “Pteropurpura” cf. tricarinatus ‌Newton, 1922], Nigeria, middle Eocene, Fig. 2C; Murexdiderrichi ‌Vincent, 1913 [?synonym of Murex ‌camerunensis ‌Oppenheim, 1904], Angola, Cameroon, late Paleocene-early Eocene; Murex ‌bicostatus ‌Deshayes, 1835, France, middle Eocene (Bartonian) (Fig. 2D, E).

Name formed by palaiós (meaning old in ancient Greek) and Chicoreus.

Biconic and middle-sized shell with Chicoreus shape. Well marked primary cords. Spire: IP (infrasutural ramp), P1 to P3 (convex part of whorl). Last whorl: IP, P1 to P6 (convex part of whorl), P6 not atrophied, ADP, MP and ABP (siphonal canal). Nodulose protovarices on early whorls. Three varices with one or two intervarices on succeeding whorls. Varices slightly winged on last whorl. No cord spine even on P1. Shoulder sinus very short. Aperture ovate with strong internal denticles within outer lip: ID, D1 to D6. Columellar lip smooth. Parietal callus and anal notch present. Aragonite microstructure.

During the Eocene several species of Paleochicoreus ‌Merle ‌n. gen., appeared in Eastern Pacific (North American coast), Western Atlantic (North American coast) and Eastern Atlantic (West Africa and Europe), which have been attributed in the literature to Pterynotus ‌Swainson, 1833 orChicoreus ‌Montfort, 1810 (Eames 1957; Vokes 1971, 1990; Merle et al. 2011 as ?Chicoreus). With Chicoreus, they share a biconic shell, medium length of siphonal canal, trivaricate morphology, nodulose protovarices, an anal notch, and a parietal callus. However, these features are also shared with the genera Naquetia ‌Jousseaume, 1880, Phyllonotus Swainson, 1833 and Chicomurex ‌Arakawa, 1964. Paleochicoreus ‌Merle ‌n. gen., differs from these three genera by having a smooth columellar lip. For example, according to Vokes (1990), Chicoreus(Phyllonotus) initialis(herein Paleochicoreus ‌initialis ‌n. comb.) (Fig. 2A, B) is most similar to members of Phyllonotus, because of developed internal denticles of the outer lip, but it lacks the derived characters of Phyllonotus (rugae and expanded parietal shield). Paleochicoreusnewtoni(Fig. 2C) possesses an expanded varical flange, like that in Paleochicoreusinitialis ‌n. comb., but this character is lacking in many Chicoreus species. Members of Chicoreus(Triplex) and C. (Rhizophorimurex) also bear similarities with those of Paleochicoreus ‌Merle ‌n. gen.,, but they lack an expanded varical flange, their internal denticles are reduced or missing, and they often develop spiny processes. Chicoreus (s.s.), which is rather similar to C. (Triplex), differs by having a labral groove spine placed between P4 and P5 (Merle et al. 2011: fig. 37A, B), lacking in Paleochicoreus ‌Merle ‌n. gen., Species such as Paleochicoreusinitialis ‌n. comb. and P.newtonirepresent ideal ancestral lineages for Chicoreus, Chicomurex or Phyllonotus (Vokes 1990) but cannot readily be assign to one of these branches. This comment is true for other species attributed here to Paleochicoreus ‌Merle ‌n. gen., Here, we figure and give a new description of the holotype of P. bicostatus ‌(Deshayes, 1835) ‌n. comb., This very rare species has not been found subsequent to its original description. Information given hereunder is based on the holotype retrieved from the collections of the École des Mines, Lyon.

Holotype (by monotypy).France• Paris Basin, Val d’Oise, Valmondois; Sables d’Auvers (biozone NP16); middle Eocene (lower Bartonian); UCBL-EM 33437 (Deshayes coll.); H: 21.5 mm (apex incomplete) (Fig. 2D, E).

Sables d’Auvers (biozone NP16), see Gély & Lorenz (1991: pl. 1), middle Eocene (lower Bartonian).

France, Paris Basin, Val d’Oise, Valmondois (see Fritel 1910: 315).

Only known from the type locality.

Teleoconch of 21.5 mm in height (apex incomplete), of 13.2 mm in width, biconic in profile, composed of 3.5 preserved whorls. Moderately high spire with convex whorls. Last whorl up to 79% of total length of teleoconch. Apical angle 63°. Spiral sculpture with well-marked primary cords. Spire with P1 and P2 cords. Last whorl: IP, P1 to P6 (P6 placed on top of siphonal canal), ADP, MP and ABP; secondary cords (adis, abis, s3-s6, ads and ms). Axial sculpture not known on early teleoconch whorls. On spire and last whorl, three varices and between them two intervarices. Intervarices strongly developed. Intersection between axial and spiral sculpture displaying small nodules on IP, P1 to P5, ADP and MP. Intervarical nodules, present and more developed on P1 and P2. Ovate aperture of 24% of diameter, up to 64% of length of last whorl. Parietal lip smooth, adherent. Columellar lip smooth, slightly erect. Anal notch deep. Parietal callus well developed. Outer lip showing internal denticles: ID stronger than D1, D2 and D3, D4, D5 and D6 stronger than D1 to D3, D6 split. Siphonal canal of 50% of apertural length. Pseudoumbilicus narrow.

No new specimens of this species have been found since its original description by Deshayes (1835). Cossmann & Pissarro (1911: pl. 36, fig. 170bis-1) attributed an eroded specimen (MNHN.F.J02588 refigured in Merle et al. 2022: pl. 147, fig. 4a, b) to this species under the name Ocenebra(Ocinebrina)bicostata ‌(Deshayes, 1835). However, re-examination of this specimen (Merle 1999) reveals that it represents Eofavartia ‌subrudis ‌(d’Orbigny, 1850). Paleochicoreus ‌bicostatus ‌(Deshayes, 1835) ‌n. comb., shares with P. ‌newtoni ‌(Eames, 1957) and P. ‌initialis ‌(Vokes, 1990) ‌n. comb., an aperture with developed internal denticles, a strong parietal callus, and a deep anal notch. P. ‌newtoni differs in having a slenderer shape and lower, less coarse intervarices. In P. ‌initialis ‌n. comb., the intervarices are also lower than in P. ‌bicostatus ‌(Deshayes, 1835) ‌n. comb., Paleochicoreus ‌packardi ‌(Dickerson, 1915) is poorly preserved (see Merle et al. 2011), but it bears slightly winged varices as do P. ‌newtoni and P. ‌initialis ‌n. comb.Paleochicoreus ‌diderrichi ‌(Vincent, 1913) displays well developed denticulation within the outer lip (see Merle et al. 2011: fig. 41.3). Paleochicoreus ‌bicostatus ‌(Deshayes, 1835) ‌n. comb., differs from these species in lacking winged varices. However, the holotype is eroded and in the early Bartonian (Auversian) from the Paris Basin, many species tend to lose their delicate ornaments (wings, spines) and thicken their shell, as is the case in another muricid, Eofavartia ‌subrudis ‌(Deshayes, 1835).

Holotype.France• Loire Basin, Loir-et-Cher, Pontlevoy; Faluns du Blésois (mammal biozone MN5b); Middle Miocene (base of the Langhian); MNHN.F.A71120 (Dollfus coll.), figured specimen in Merle et al. (2022: pl. 3, fig. 4), H: 14 mm (Figs 3A, B; 25A).

Paratypes. France• 1 spm; same data as for the holotype; MNHN.F.A71122, figured specimen in Merle et al. (2022: pl. 3, fig. 5)•11 spm; idem;MNHN.F.A77742 (Dollfus coll.)•1 spm; idem; MNHN.F.A77743 (Dollfus coll., Figs 3C, D; 25B)•spm; idem; MNHN.F.B26846 (MNHN coll.)•5 spm; idem; MNHN.F.A77744 (Lhomme coll.);•2 spm; LoireBasin, Indre-et-Loire, Paulmy (Pauvrelay); idem; MNHN.F.A77745 (Faullummel coll.).

The name pontileviensis refers to the type locality: Pontlevoy.

Faluns du Blésois (mammal biozone MN5b), see Ginsburg (2000), Middle Miocene (base of the Langhian).

France, Loire Basin, Loir-et-Cher, Pontlevoy (see Macaire et al. 2020).

Loire Basin (France), Faluns du Blésois, Middle Miocene (Langhian).

Paucispiral and rounded protoconch of 1.5 whorls. Teleoconch up to 14 mm in height, up to 6.5 mm in width, biconic in profile, composed of six whorls. Moderately high spire with subcarinate whorls. Last whorl up to 67.7% of total length. Apical angle 42°. Spiral sculpture with marked primary cords. First whorl: appearance of P1 and P2 (P2 more developed than P1) at beginning of whorl and appearance of IP on second half of whorl; second to fifth-sixth whorl: no change, but sometimes s3 placed slightly above suture. Sixth whorl: IP (sutural ramp), P1 to P5 (convex part of whorl, P6 atrophied (on siphonal canal), ADP and MP well developed, ABP poorly developed. Secondary cords s1, s4, s5 and s6 appear between end of fourth whorl and fifth whorl; s2 and s3 appear earlier. On first whorl: 14 varices; from second to fourth whorl: 10-12 varices; from fifth to sixth whorl: 10-11 varices. On spire axial ornamentation forming small nodules placed on IP, P1 and P2. P1 and P2 nodules more developed than IP. On convex part of last whorl, P1 to P4 nodules more or less equally developed; P5 slightly more developed; on siphonal canal, P6 not atrophied, ADP and MP more developed than ABP when present. Ovate aperture up to 42% of diameter and up to 70% of length of last whorl (including siphonal canal). Columellar lip narrow, entirely adherent, bearing two weak tubercles anteriorly. No anal sulcus, no parietal callus. Outer lip with strong internal denticles. Series of denticles including: ID, D2 to D5. D1 missing. D2 stronger than other denticles. Outer lip not crenulate. Siphonal canal open, up to 33% of apertural length and slightly dorsally recurved. Pseudo-umbilicus narrow.

This species is comparable to Muricopsis ‌crassicostata ‌(Benoist, 1880) from the late Oligocene/Early Miocene of Aquitaine Basin (France) and M. ‌dujardini ‌Peyrot, 1938 from the Middle Miocene of Loire Basin (France). Muricopsis ‌crassicostata clearly differs from M. ‌pontileviensis ‌n. sp. in being larger (19-23 mm for 6-7 whorls vs 12-14 for 5.5-6 whorls) and having a multispiral protoconch of three convex whorls (for more comparative details, see Merle 1999: 348, fig. 115). Muricopsis ‌dujardini and M. ‌pontileviensis ‌n. sp. both occur at Pontlevoy. Specimens of M. ‌dujardini are larger (19-24 mm for 6-6.5 whorls), but they are mainly distinguishing from M. ‌pontileviensis ‌n. sp. by having short spines on P1 to P5 in which P3 is atrophied and by having stronger denticles within the outer lip. Muricopsis ‌pontileviensis ‌n. sp. never bears spines. Merle (1999) wrongly attributed studied specimens of M. ‌pontileviensis ‌n. sp. to M. ‌dujardini, however, he demonstrated that they display a paedomorphic shape (particularly because of the post-displacement of the secondary cords s1, s4 and s6) compared to larger shells as M. ‌crassicostata.

Holotype. France• Rennes Basin, Ille-et-Vilaine, Saint-Jacques-de-la-Lande; Yellow bed of the quarry; Late Miocene (lower Messinian); MNHN.F.A77746 (Néraudeau’s sample), H: 8.5 mm (Figs 3E, F; 25C).

Paratypes. France• 1 spm; same as for the holotype; MNHN.F.A77747 (Néraudeau sample, Figs 3G, H; 25D)•12 spm; idem; MNHN.F.A90516 (Néraudeau’s sample).

Dedicated to Didier Néraudeau who collected and provided the material.

Yellow bed of the quarry, Late Miocene (lower Messinian), see Néraudeau et al. (2002).

France, Rennes Basin, Ille-et-Vilaine, Saint-Jacques-de-la-Lande (Lilion sand pit).

Only known from the type locality.

Protoconch not preserved. Teleoconch up to 8.5 mm in height, up to 4 mm in width, biconic in profile, composed of five whorls. Moderately high spire with subcarinate whorls. Last whorl up to 69.6% of total length. Apical angle 50°. Spiral sculpture with coarse primary cords. First whorl eroded; second whorl appearance of IP; P2 more developed than P1; second to fourth whorl: no change (IP, P1 and P2). Fifth whorl (last whorl): IP, P1 to P5, P6 atrophied (on siphonal canal), ADP and MP well developed, ABP poorly developed. Secondary cords s2 and s3. On second whorl: 12-13 varices; on fourth whorl: 10-11; on fifth whorl: 9-10 varices. On spire, axial ornamentation forming marked nodules on IP, P1 and P2. P1 and P2 nodules more developed than IP nodule. On convex part of last whorl, P1 to P5 acute nodules more or less equally developed; on siphonal canal, only ADP and MP displaying nodules. Ovate aperture up to 44% of diameter and up to 72% of length of last whorl (including siphonal canal). Columellar lip narrow, entirely adherent, bearing two tubercles anteriorly. No anal sulcus; no parietal callus. Outer lip with strong internal denticles. Series of denticles including ID, D2 to D5. D1 missing. D2 stronger. Outer lip not crenulate. Siphonal canal open, up to 32% of apertural length, slightly dorsally recurved. Pseudo-umbilicus narrow.

Like Muricopsis ‌pontileviensis ‌n. sp., M. ‌neraudeaui ‌n. sp. is another very small species for the genus and is comparable to the Langhian M. ‌pontileviensis ‌n. sp. in size. However, M. ‌neraudeaui ‌n. sp. is even smaller (8.5 mm for 5 whorls instead 12-14 mm for 5.5-6 whorls). Muricopsis ‌neraudeaui ‌n. sp. displays a stronger sculpture formed by acute nodules on IP, P1 to P5 and ADP, MP), whereas M. ‌pontileviensis ‌n. sp. bears only a swelling on the varices. In addition, the secondary cords are less numerous in M. ‌neraudeaui ‌n. sp. (only s2 and s3 instead s1, s2, s3, s4, s5 and s6 in M. ‌pontileviensis ‌n. sp.). In spite of its very small size, M. ‌neraudeaui ‌n. sp. bears strong columellar denticles. This is the first record of very small Muricopsis species in fossil record. Assemblages containing very small species of Muricopsis are only documented in the Recent faunas. The best examples come from the islands of São Tomé and Martinique which contain several very small Muricopsis species (Houart 1996; Garrigues & Lamy 2019 respectively).

Holotype.The Netherlands• North Sea Basin, Miste, Winterswijk; Miste bed; Middle Miocene (middle Langhian); RGM.225979 (figured specimen in Janssen [1984: pl. 59, fig. 7]), H: 12.5 mm (Figs 4A, B; 25E).

Paratype.The Netherlands• 1 spm; same as for the holotype; MNHN.F.A77748 (Ledon coll., Figs 4C, D; 25F).

Dedicated to Arie W. Janssen for his paleontogical and malacological work.

Miste bed, Middle Miocene (middle Langhian), see Janssen (1984) and Schneider & Heissig (2005).

The Netherlands, North Sea Basin, Miste, Winterswijk (see Janssen 1984).

Only known from the type locality.

Protoconch not preserved. Delicate teleoconch up to 12.5 mm in height, up to 7.5 mm in width, composed of five whorls. Moderately high spire with subangular whorls. Last whorl up to 72% of total length. Apical angle 72° (including spines) and 48° (excluding spines). Spiral sculpture with strong primary cords. From first to fourth whorl: P1 and P2; on fourth whorl, appearance of IP. Fifth whorl: IP and abis, P1 to P5, P6 atrophied (on siphonal canal), ADP and MP present, ABP missing. Surface of P1, P2 and P3 bearing deep spiral grooves. Second whorl: 8-9 varices; from third to fourth whorl: 8 varices; fifth whorl: 6-7 varices. Adapertural face of varices with some imbricated growth lamellae. Axial ornamentation forming short spines on P1 to P5, ADP and MP. Axial microsculpture with small erect lamellae. Suboval aperture up to 29% of diameter and up to 71% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, erect anteriorly. Outer lip lacking denticles. Siphonal canal open, up to 42% of apertural length and slightly dorsally recurved. Pseudo-umbilicus rather wide.

This rare species from the Langhian of Miste (North Sea) was identified as Favartia ‌collega ‌(Boettger, 1906) by Janssen (1984). The holotype of F. ‌collega (SMF XII2258a, figured in Merle et al. 2022: pl. 58, fig. 2) comes from the Langhian of Costeiu de Sus (Romania, Paratethys Sea). It differs from F. ‌jansseni ‌n. sp. by having a slenderer shell, a rounded shoulder sinus in the aperture, and weak internal denticles within the outer lip (D1 to D5). Secondary cords are poorly developed, but the holotype of F. ‌collega is a juvenile specimen of only 7.5 mm height. Favartia ‌collega and F. ‌czjzeki ‌ (Hoërnes & Auinger, 1885) from the Langhian of Austria (Paratethys Sea) and Hungary (Kovács et al. 2018) share a slender shell, a rounded shoulder sinus, and a multispiral protoconch (Kovács et al. 2018). The shells of F. ‌collega and F. ‌czjzeki are very similar, and therefore, F. ‌collega is considered to be a junior synonym of F. ‌czjzeki herein. The syntype of F. ‌czjzeki (NHMW 1855/0045/008 figured in Merle et al. 2022: pl. 58, fig. 1) and other adult specimens of this species display the secondary cords (s1, s2, s3 and s5) and weak internal denticles missing in F. ‌jansseni ‌n. sp.

Holotype. France• Loire Basin, Loir-et-Cher, Pontlevoy; Faluns du Blésois (mammal biozone MN5b); Middle Miocene (base of the Langhian); MNHN.F.A33456 (Dollfus coll.) figured specimen in Merle et al. (2011: pl. 173, fig. 1a-b), H: 8 mm (Figs 4E, F; 25G).

Paratype. France• 1 spm; same as for the holotype; MNHN.F.A33457 (Dollfus coll.) figured specimen in Merle et al. (2011: pl. 173, fig. 2a, b) (Figs 4G, H; 25H).

From the Latin adjective ligerianus meaning Loire.

Faluns du Blésois (mammal biozone MN5b), see Ginsburg (2000), Middle Miocene (base of the Langhian).

France, Loire Basin, Loir-et-Cher, Pontlevoy (see Macaire et al. 2020).

Only known from the type locality.

Bulbous protoconch of 1.5 whorls. Teleoconch of up to 8-9 mm in height, up to 4 mm in width, composed of five elongated whorls. Spire high, weakly convex whorls. Last whorl up to 67% of total height. Apical angle 45°. Spiral sculpture with moderately marked primary cords. First to second whorl: no cord. Third whorl: appearance of P2 on base of whorl. Fourth whorl: appearance of IP and P1 (weak), presence of P2 and P3. Fifth whorl: IP, P1 to P5 on convex part of whorl, siphonal canal smooth. From first to second whorl, six lamellose protovarices extending between sutures. On third whorl, 6-7 protovarices. From fourth to fifth whorl, two thickened varices placed at periphery, 180° to each other, with two intervarices intercalated. Bivaricate morphology giving shell laterally compressed appearance. No cord spine, but only flat nodules at intersection between axial sculpture and major cords (P1 to P5). Aperture ovate, up 35% of diameter, up to 67% of height of last whorl (including siphonal canal). Columellar lip smooth poorly erect anteriorly. Outer lip with very weak internal denticles. Series of internal denticles including ID, D1 to D4. D5 and D6 not developed. Pseudo-umbilicus narrow. Siphonal canal up to 35% of apertural lenght. Remains of intritacalx near varices.

All other members of the subgenera of Dermomurex [Dermomurex (s.s.), Takia ‌Kuroda, 1953 and Viator ‌E. H. Vokes, 1974] are characterized by a series of internal denticles containing ID, D1 to D5 or D6, whilst D5 and D6 are missing in the two known species of this subgenus: D. (Gracilimurex)bakeri ‌(Hertlein & Strong, 1951), Recent, Eastern Pacific (Gulf of California) and D.(G.)elisabethae ‌(McGinty, 1940), Pleistocene to Recent, Western Atlantic (Florida, Mexico). Nevertheless, only D.(Gracilimurex)ligerianus ‌n. sp. bears a poorly developed D5, but this character state, restricted in this old species of D. (Gracilimurex), could be interpreted as the retention of the primitive state, the derived state being the loss of this denticle. The bivaricate morphology found in D. (Gracilimurex) is unique in Dermomurex and can be compared to Aspella. However, it appears early during ontogeny in Aspella, whereas it appears later (on the penultimate or the last whorl) in D. (Gracilimurex). According to Vokes (1992: 72), the shell ornamentation of D. (Gracilimurex) is more closely akin to that of Dermomurex (s.s.), especially the group of the type species. The oldest record of D. (Gracilimurex) was D.(Gracilimurex)elisabethae, found in the Pleistocene (Calabrian, Bermont Formation) of Florida. Therefore, this new species from the Middle Miocene of Loire Basin predates the previous oldest record by 12 million years and expand the geographic range of the subgenus to the Eastern Atlantic.

Holotype. France• Aquitaine Basin, Pyrénées-Atlantiques, Gan (Tuilerie); Uppermost part of the Marnes de Gan only (top biozone N12/base biozone NP13); Early Eocene (Ypresian); MNHN.F.A90520 (Varone coll.), H: 17 mm (Figs 5A, B; 25I).

Paratype. France• 1 spm; same as for the holotype; MNHN.F.R61441 (Merle coll., figured specimen of Merle (1990: pl. 3, fig. 13)•1 spm; idem; MNHN.F.A90518 (Merle coll., with complete siphonal canal, Figs 5C; 25J)•1 spm; idem; MNHN.F.A90517 (Merle coll. with incomplete siphonal canal; Figs 5D; 25K)•12 spm; idem; MNHN.F.A90519 (Merle coll.)•17 spm; idem; MNHN.F.A72383 (Varone coll.).

4 spm; idem; MNHN.F.A90521 (Combault coll.).

From the type locality, Gan (Pyrénées-Atlantiques).

Uppermost part of the Marnes de Gan only (top biozone N12/base biozone NP13), see Nolf et al. (2002: 173, fig. 4), Early Eocene (Ypresian).

France, Aquitaine Basin, Pyrénées-Atlantiques, Gan (Tuilerie) (Merle 1986: 8; Nolf et al. 2002: 185, 186).

Only known from the type locality.

Bulbous protoconch of 1.5-2 whorls (see Merle 1990: pl. 4, fig. 12). Teleoconch up to 17 mm (canal siphonal not complete) in height, up to 11 mm in width, biconic in profile, composed of five whorls. Moderately high spire. Last whorl representing 72% of total height. Spiral angle 56°. Suture weakly impressed. Axial sculpture consisting in four slightly foliate varices per whorl. On spire, anal tube (P1) and very short spine (P2). Anal tube forming angle of approximately 60-65° with axis of shell. Anal tube (P1) totally integrated within varices, eliminating shoulder. Small callus at base of anal tube. On convex part of last whorl and abapically to anal tube four spinelets (P2, s2, P3, P4 and P5). Intervarical spaces usually smooth, but with P3-P5 cords on holotype. Rounded aperture up to 25% of diameter, up to 74% of height of last whorl (including siphonal canal). Edge, erect, smooth, peristome complete. Siphonal canal closed, usually broken, occupying 60% of apertural lenght (complete siphonal canal longer, see Fig. 5C).

Merle (1990: 169) attributed this material collected at Gan to Typhis ‌parisiensis ‌d’Orbigny, 1850 and to the genus Trubatsa ‌Dall, 1889. The generic attribution is correct and the distinction between Siphonochelus ‌Jousseaume, 1880 and Trubatsa is now validated by molecular data (Houart et al. 2021). Nevertheless, after further comparison, specific attribution of the Gan material to this species incorrect. Firstly, the protoconchs are different, that of T. ‌ganensis ‌n. sp. is paucispiral (Merle 1990: pl. 4, fig. 12) and that of T. ‌parisiensis is multispiral (see juvenile specimen from Barton illustrated by A. Morton (2023; www.dmap.co.uk/fossils). In addition, the Bartonian specimens are more sculptured than the specimens from Gan (Fig. 5E, F). They display stronger spines (P2 to P5) and P6, spiral cords (P2 to P5) in the intervarical spaces and a partition recalling that of T. ‌gaasensis (Tournouër in Benoist, 1880) from the Rupelian of Aquitaine Basin. Trubatsa ‌parisiensis occur in the Bartonian from England and in the Priabonian from Germany (von Koenen 1889). Merle (1985) also suggested that Typhis ‌sinuosus ‌Cossmann, 1902 from the Bartonian of Bois-Gouët (France, Loire-Atlantique) was a junior subjective synonym of T. ‌parisiensis. According to Merle (1985), the shape of the varices of the type specimen (Dumas coll.) of T. ‌sinuosus recalls that of Trubatsa. However, it is a juvenile (H: 8 mm) and the varices of the juveniles of Typhina ‌rutoti ‌Cossmann, 1882 are also similar. Moreover, the type specimen of T. ‌sinuosus is worn, lacks a protoconch and finally lacks diagnostic characters. Thus, we consider T. ‌sinuosus as a nomen dubium.

Holotype. France• Paris Basin, Oise, Cauvigny (Châteaurouge); Calcaire grossier moyen (biozone NP15); middle Eocene (middle Lutetian); MNHN.F.A25590 (Pacaud coll.), H: 17 mm (Figs 5G, H; 25L).

From Calviniacus, the Latin name of the type locality Cauvigny (Oise, France).

Calcaire grossier moyen (biozone NP15), see Gély & Lorenz (1991: pl. 1); middle Eocene (middle Lutetian).

France, Paris Basin, Oise, Cauvigny (Châteaurouge), see Fritel 1910: 85).

Only known from the type locality.

Bulbous protoconch of two whorls; pointed apex. Teleoconch up to 17.8 mm in height, up to 9.2 mm in width, biconic in profile, composed of six whorls. High spire. Last whorl representing up to 70% total height. Spiral angle 62°. Suture weakly impressed. Axial sculpture consisting of four slightly spiny varices per whorl. On spire, anal tube (P1) and short spine (P2). Anal tube forming angle of approximately 45° with shell axis. Anal tube (P1) totally integrated in varices eliminating shoulder. Short P2 spine on first whorls. No callus at base of anal tube. On convex part of last whorl and abapically to anal tube, five spinelets (P2, P3, P4, P5 and P6). Intervarical spaces with weak P2 to P5 cords. Ovate aperture up to 30% of diameter, up to 74% of height of last whorl (including siphonal canal). Edge, erect, smooth. peristome complete. Siphonal canal closed, occupying 47% of apertural lenght.

Merle & Pacaud (2019: pl. 3, fig. 4) attributed a specimen (MNHN.F.A25590) from the middle Lutetian of Cauvigny (Paris Basin) to T. ‌parisiensis ‌(d’Orbigny, 1850). Like T. ‌ganensis ‌n. sp. it has a paucispiral protoconch and lacks partition. Therefore, it can easily be distinguished from T. ‌parisiensis from the Hampshire Basin which has a multispiral protoconch. Trubatsa ‌calviniacensis ‌n. sp. differs from T. ‌ganensis ‌n. sp. in being higher spired, the primary cord P6 on the last whorl, weak primary cords P2 to P4 in the intervarical spaces, and by having a more ovate aperture. P2 spine is also more marked on the first teleoconch whorls. Although we have only one specimen, the number of shell differences with the Gan specimens and its occurrence in a strongly different paleoenvironment (deep water at Gan (Merle 1985; Merle & Roux 2018) in Aquitaine Basin, versus shallow water in the Paris Basin (Gély 2008) and also quite different age, highly suggest that it represents a different species.

Murex ‌latifolius ‌Bellardi, 1873, by subsequent designation (Vokes 1964: 14). Middle Miocene: Italy.

Historically, fossil and Recent members of Timbellus have usually been attributed to the genus Pterynotus ‌Swainson, 1833 (e.g., Cossmann 1889; Cossmann & Pissarro 1911; Glibert 1963; Vokes 1970) in the subfamily Muricinae. Based on molecular data, Barco et al. (2010, 2012) studied the phylogenetic relationships of Pterynotus ‌Swainson, 1833 represented byP. ‌elongatus ‌(Lightfoot, 1786) and P. ‌fulgens ‌Houart, 1988 and Pterymarchia ‌Houart, 1995 represented by P. ‌martinetana ‌(Röding, 1798). These authors demonstrated firstly, that Pterynotus and Pterymarchia are not phylogenetically close to P. ‌fulgens ‌Houart, 1988 and secondly, none of these taxa are included within the clade of the Muricinae. According to Merle et al. (2011), Pterynotus ‌fulgens is closely related to the type species of Timbellus and is now attributed to that genus. Thus, Merle et al. (2011) and Houart (2018) excluded Timbellus from the Muricinae, but did not give a subfamilial attribution. Russini et al. (2023: 866) place Timbellus in incertae sedis and considered it likely to represent an independent lineage worthy of subfamilial rank.

In order to delineate different lineages included in Pterynotus (s.s.), Harasewych & Jensen (1979) and more recently Vokes (1992) defined five “species groups” for the Western Atlantic Region. With the exception of the typical group of Pterynotus [species group of Pterynotus ‌pinnatus ‌(Swainson, 1822)], the four other species groups are members of the genus Timbellus ‌(Merle et al. 2011). Merle et al. (2011: 130) added a fifth group to Timbellus: the species group of T.tripteroides ‌(Lamarck, 1822). Members of that group display a strong development of the internal denticles within the outer lip and their shell shape is narrower than those belonging to the group of T. ‌crenulatus ‌(Röding, 1798). Fine growing lamellae forming a scabrous surface are usually missing and only reported in T. ‌capitaneus ‌Pacaud, Ledon & Goret, 2017. The following new species described herein, T. ‌magnificus ‌n. sp., T. ‌occidentalis ‌n. sp., T. ‌calciacus ‌n. sp., T. ‌longicanalis ‌n. sp. and T. ‌magnei n. name (for Murex ‌trigonus ‌Rouault, 1850, non Murex trigonus Gmelin, 1791) display features allowing a placement in this species group. Several of these new species (T. ‌occidentalis ‌n. sp., T. ‌calciacus ‌n. sp., T. ‌longicanalis ‌n. sp. and T. ‌magnei n. name) are superficially similar to, and mistaken for, T. ‌tripteroides in the past. Moreover, the type material of T. ‌tripteroides is lost, as it is missing in the Lamarck’s collections in Geneva and Paris. In order to avoid future taxonomic mistakes, we designate a neotype herein (MNHN.F.A90546, coll. of the Club géologique d’Île-de-France, Fig. 6B, C). The type material of Lamarck was collected at Grignon (middle Lutetian) and the neotype is selected from the same type locality (precisely Falunière of Grignon; Calcaire à Orbitolites complanatus Formation, biozone NP15, see Gély & Lorenz 1991). The neotype was chosen to closely represent the specimen illustrated by Lamarck (1805: pl. 3, fig. 4; Fig. 6A). It displays rounded wings, P1 spine is included in the wings and is not protruding. This morphology of the anterior part of the wings (Fig. 6D) differs from that of the Murex ‌tripteroides illustrated by Deshayes (1835: pl. 82, figs 1, 2; Fig. 6E, F) and Cossmann & Pissarro (1911: pl. 34, 169-1), which display a well individualized P1 spine (Fig. 6G, H). We examined numerous specimens from different localities in the Paris Basin and found transitional forms between the two morphotypes. Thus, we consider this difference to fall within the intraspecific variation.

Holotype. France•Paris Basin, Oise, Baron (Sablière Heudebert); Sables d’Auvers (biozone NP16); middle Eocene (lower Bartonian, Auversian); MNHN.F.A90522 (Pacaud coll.), H: 58.5 mm (Figs 7A, B; 25M).

Paratypes.France• 1 spm; same as for the holotype; early Bartonian; MNHN.F.A90523 (Pacaud coll.) (Figs 7E, F; 25O)•3 spm; idem; MNHN.F.A90524 (Faullummel coll.)•1 spm Paris Basin, Oise, Saint-Vaast-lès-Mello (Barisseuse); idem; MNHN.F.A90525 (Schtrock coll.) (Figs 7C, D; 25N)•1 spm; Paris Basin, Aisne, Montreuil-aux-Lions; idem; MNHN.F.A73852 (Pons coll.)•1 spm; Paris Basin, Seine-et-Marne, Isle-les-Meldeuses; idem; MNHN.F.A90526 (Faullummel coll.)•1 spm; Paris Basin, Val d’Oise, Auvers-sur-Oise (Bois-le-Roi); idem; MNHN.F.A90527 (ex galerie de Zoologie coll.).

France • 1 spm; Paris Basin, Aisne, Bézu-le-Guéry; Sables d’Auvers (biozone NP16); early Bartonian; MNHN.F.A90528 (Faullummel coll.)•1 spm; Paris Basin, Aisne, Baron (Sablière Heudebert); idem; MNHN.F.A90529 (Merle coll.)•1 spm; Paris Basin, Seine-et-Marne, Le Limon; idem; MNHN.F.A90530 (Pacaud coll.).

From the Latin magnificus meaning magnificent.

Sables d’Auvers (biozone NP16), see Gély & Lorenz (1991, pl. 1), middle Eocene (lower Bartonian, Auversian).

France, Paris Basin, Oise, Baron (Sablière Heudebert), see Wyns (1980).

Paris Basin (France), Sables d’Auvers, middle Eocene (lower Bartonian, Auversian).

Unknown not preserved. Teleoconch up to 58.5 mm in height, up to 33 mm in width, subfusiform in profile, composed of seven whorls. Moderately high spire. Last whorl up to 85% of total length. Apical angle 71°. Spiral sculpture with moderately marked primary cords on varices. From first to fourth whorl, no spiral sculpture. Fifth whorl: appearance of weak cords corresponding to P1, P2, P3, s1 and s2; primary and secondary cords of same thickness. Sixth whorl: increase of thickness of cords. Seventh whorl: P1 to P6, s1 to s5, P6 weak; ADP, MP and ABP on siphonal canal. Axial sculpture eroded on first whorl. On second whorl, eight protovarices. On third whorl: appearance of varices; three varices and one intervarix between. From fourth to last whorl no change; intervarices low. Higher relief intervarices around on P2. Varices sublamellose on early whorls, thickened on last whorls. Appearance of short P1 spine on fourth whorl. On last whorl, short P1 spine oriented adapically; no other cord spine. Ovate aperture up to 36% of diameter and up to 70% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, slightly erect anteriorly and forming very flat curved inductura at base. Parietal lip slightly adherent. Outer lip with small denticles, ID, D1, D7 simple, D2-D6 bifid or trifid for D4. Siphonal canal narrow, open, bent, slightly dorsally recurved, up to 43% of apertural length. Pseudoumbilicus narrow.

With a length exceeding 55 mm, it is one of the largest Eocene Timbellus with T. ‌capitaneus ‌Pacaud, Goret & Ledon, 2017 which can exceeed a length of 65 mm. The spiral cords of T. ‌magnificus ‌n. sp. are weak in juvenile specimens of four whorls, whereas they are already well developed and well marked in juvenile specimens of T. ‌capitaneus at the same growth stage. In addition, T. ‌magnificus ‌n. sp. bears a small and very short P1 spine oriented adapically, whereas that of T. ‌capitaneus is strong and abaxially oriented. Moreover, T. ‌capitaneus and T. ‌magnificus ‌n. sp. share a series of internal denticles including D1 to D6, but all internal denticles of T. ‌magnificus ‌n. sp. are bifid and more or less of the same size.

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; LoireBasin, 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).

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

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

France, Normandy, Manche, Fresville, see APGN (2024).

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).

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.

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. 8A-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.

Holotype. France• Paris Basin, Val d’Oise, Chaussy (Les Garennes); Falun de Chaussy (biozone NP15); middle Eocene (middle Lutetian); MNHN.F.A90536 (Merle coll.), H: 8 mm (Figs 10A, B; 26C).

Paratypes. France• 3 spm; same as for the holotype; MNHN.F.A90537 (Figs 10C, D; 26D, H: 12 mm), MNHN.F.A90538, MNHN.F.A90539 (Merle coll.)•1 spm; idem; MNHN.F.R61443 (Merle coll.), figured specimen of Merle (1990: pl. 4, fig. 3)•1 spm; Paris Basin, Yvelines, Beynes (Ferme de l’Orme); middle Lutetian; MNHN.F.A90535 (Merle coll.).

Calciacus (Latin adjective): from the ancient name of the village of Chaussy given in 690. Name given in apposition.

Falun de Chaussy (biozone NP15), see Gély & Lorenz (1991: pl. 1), middle Eocene (middle Lutetian).

France, Paris Basin, Val d’Oise, Chaussy (Les Garennes), see Fritel (1910: 88).

Faluns de Chaussy and Calcaire à Orbitolites at La Ferme de l’Orme (biozone NP15), Paris Basin (France), middle Eocene (middle Lutetian only).

Bulbous protoconch of 1.2 whorls. Teleoconch up to 12 mm in height, up to 5.5 mm in width, biconic in profile, composed of four whorls. Moderately high spire. Last whorl up to 76% of total length. Apical angle 71° including winged varices, up to 55° excluding them. Spiral sculpture with poorly marked primary cords. First whorl: appearance of very weak P1, P2 and P3. Second whorl: weak P1 to P3. Third whorl: appearance of weak s1, s2 and threads covering base of whorl. Fourth whorl: P1 to P6 weak, ADP on siphonal canal, secondary cords undistinguishable from threads; threads covering entire surface of shell. On first whorl, axial sculpture with 8-9 lamellose protovarices. On second whorl: appearance of varices (three varices and one low intervarix between two varices). From third to last whorl, numerous growing microlamellae covering surface of shell and forming delicate, reticulate sculpture covering surface of shell. Intervarical relief very low, but slightly higher on P1-P2. Varices well winged on last two whorls. Appearance of short P1 spine on second whorl. On last whorls, axial ornamentation forming winged varices, rounded in outline, without evidence of spines, even on P1. Subtriangular aperture up to 33% of diameter and up to 74% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, slightly erect anteriorly and forming a curved inductura at base. Parietal lip slightly adherent. Outer lip with very weak denticles including D1 to D5. Siphonal canal open, up to 45% of apertural length. Pseudo-umbilicus narrow.

This small species was never distinguished from its congeners in the Paris Basin, probably mistaken for juveniles of T. tripteroides ‌(Lamarck, 1822), with which it co-occurs at two localities: Chaussy and La Ferme de l’Orme. With a maximum height of 12 mm, T. ‌calciacus ‌n. sp. is a very small Timbellus. Timbellus ‌calciacus ‌n. sp. differs from juveniles of T. tripteroides in having the aperture completely formed and by having a peculiar reticulate microsculpture (see Fig. 6C, D). This last character lacks in specimens of T. tripteroides at any growth stage. Specimens of T. ‌occidentalis ‌n. sp. from the upper Lutetian of Normandy and the early Bartonian of Loire-Atlantique share with T. ‌calciacus ‌n. sp. a small size, but T. ‌occidentalis ‌n. sp. is double the size (H: 26.6 mm vs 12 mm). As with T. tripteroides, T. ‌occidentalis ‌n. sp. lacks the peculiar microsculpture found in T. ‌calciacus ‌n. sp.

Holotype. France• Normandy, Manche, Fresville; Calcaire de Fresville; middle Eocene (uppermost Lutetian); MNHN.F.A24101 (Le Marchand coll.), H: 35 mm (Figs 11A, B; 26E).

Paratype. France• 1 spm; same as for the holotype; MNHN.F.A24100 (Le Marchand coll.) (Figs 11C, D; 26F).

From the Latin adjective longus meaning elongated and canalis meaning canal, refecting the elongated siphonal canal of this species.

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

France, Normandy, Manche, Fresville, see APGN (2024).

Normandy (Calcaire de Fresville); middle Eocene (uppermost Lutetian).

Protoconch not preserved. Teleoconch up to 35 mm in height, up to 14 mm in width, fusiform in profile, composed of six whorls. Moderately high spire. Last whorl up to 77% of total length. Apical angle 41° excluding spine and 61° including spines. Spiral sculpture with moderately marked primary cords on varices. First and second whorl eroded. Third whorl: appearance of weak s1 and P2 on intervarices. Fourth whorl: appearance of weak P1 cord. Fifth whorl: no change. Sixth whorl: P1-P4 marked, P5-P6 weak, s1, s2, s6, P6 not atrophied, s1-s2, s6 weak; ADP, MP and ABP weak. Axial sculpture eroded on two early whorls. From third last whorl: three varices and one intervarix intercalated. Higher relief of intervarices on P2. Varices lamellose, slightly winged. Appearance of P1 spine on third whorl, no other spines adapically. On last whorl P1 spine short, straight, slightly bent adaxially. On last whorl growth lamellae erect giving scabrous aspect to shell surface. Ovate aperture up to 29% of diameter and up to 78% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, slightly erect anteriorly, forming very slightly curved inductura at base. Parietal lip adherent. Outer lip with simple, weak denticles including ID, D1 to D6. Siphonal canal narrow, open, straight, up to 59% of apertural length. Pseudoumbilicus narrow.

This species was previously identified as Timbellus ‌fusoides ‌(Deshayes, 1865) by Merle et al. (2011: pl. 101, fig. 5a, b). Nevertheless, attribution to Murex ‌fusoides is problematic, as the type material on which that Bartonian species is based is poorly preserved. It corresponds to two syntypes housed in the collection of the University Lyon (no. UCBL-EM 33383-84; Fig. 11E). The largest is a strongly eroded specimen with most of its axial sculpture, particularly its winged varices lost. This loss of sculpture gives the shell an artificially fusiform shape. The second specimen is smaller and also strongly eroded, but P1 spine is rather turned adapically as in some Timbellus ‌tripteroides ‌(Lamarck, 1822) and its aperture resembles that of T. ‌tripteroides. We designate herein this smaller specimen (UCBL-EM 33384; Fig. 11F, G) as the lectotype of M. ‌fusoides. Therefore, M. ‌fusoides can be regarded as a possible junior synonym of T. ‌tripteroides or a nomen dubium as it is impossible to clearly identify a species based on the type material. Moreover, Bartonian specimens from the Paris Basin identified as T. fusoides are present in several collections housed in the MNHN (e.g., Faullummel, Pons and Schtrock colls). Examination of these specimens shows them to represent eroded T. ‌tripteroides (probably reworked from the Lutetian) or to large and eroded Pterochelus ‌contabulatus ‌(Lamarck, 1803). The type material of T. ‌longicanalis ‌n. sp. comes exclusively from the upper Lutetian of Normandy. It is very well preserved and its delicate and winged varices are not eroded. Thus, its fusoid shape is not due to a post-mortem alteration. The species differs from T. ‌tripteroides by having more staged and more quadrate whorls, more adaxially oriented P1 spines, a straighter siphonal canal, a less rounded aperture, and by having weaker internal denticles and spiral cords.

Neotype.France• Aquitaine Basin, Pyrénées-Atlantiques, Bos d’Arros; uppermost part of the Marnes de Gan only (top biozone N12/base biozone NP13); Early Eocene (Ypresian); BMNH 69802 (Seaman coll.) designated by Merle (1990: 158, pl. 2, fig. 14) (Fig. 12A).

France • 1 spm Aquitaine Basin, Pyrénées-Atlantiques, Gan (Tuilerie); uppermost part of the Marnes de Gan only (top biozone N12/base biozone NP13); Early Eocene (Ypresian, uppermost part of the Marnes de Gan); MNHN.F.A32506 (Merle coll.), figured specimen of Merle et al. (2011: pl. 100, fig. 3)•1 spm; idem; MNHN.F.A32508 (Merle coll.), figured specimen of Merle et al. (2011: pl. 100, fig. 1) (Figs 12C, D; 26G)•1 spm; idem; MNHN.F.R61437, Merle coll., figured specimen of Merle (1990: pl. 3, fig. 1)•1 spm; idem; MNHN.F.A24115 (Merle coll.)•2 spm; idem; MNHN.F.A72443 (Varone coll.) (Figs 12E, F; 26I)•35 spm; idem; MNHN.F.A24114 (Merle coll.)•1 spm (juvenile with protoconch); idem; MNHN.F.A90545 (Merle coll.) (Figs 12B; 26H).

Uppermost part of the Marnes de Gan only (top biozone N12/base biozone NP13), see Nolf et al. (2002: 173, fig. 4), Early Eocene (Ypresian).

France, Aquitaine Basin, Pyrénées-Atlantiques, Bos d’Arros (Nolf et al. 2002: 185).

Known from the type locality and Gan (tuilerie), Aquitaine Basin (France), early Eocene (Ypresian).

Smooth, multispiral protoconch of 3.5 whorls (Figs 12B; 26H). Sinusigeral scar well marked. Teleoconch up to 46 mm in height, up to 39 mm in width, subfusiform in profile, composed of eight whorls. Moderately high spire. Last whorl up to 70% of total length. Apical angle 53° excluding wings, 70° including wings. Spiral sculpture with marked primary cords on varices and intevarices. First whorl: no cord. Second whorl: appearance of weak P1, P2 and s1. Third whorl: P1, P2, s1 well marked; appearance of s2 weak. Fourth whorl: s2 well marked. From fifth to seventh whorl: no change or appearance of threads. Eighth whorl: P1 to P6, s1, s2, s3 (well developed), P6; ADP, MP weak or missing on siphonal canal. First whorl: around ten protovarices. Second whorl: appearance of varices, three varices and two intervarices between. From third to last whorl, three to five low intervarices. Higher relief of intervarices on P2. Varices lamellose and strongly winged. Appearance of short P1 spine on second whorl. P1 spine short, almost included in wing. No other cord spines. Ovate aperture up to 22% of diameter and up to 71% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, erect anteriorly, forming curved inductura at base. Parietal lip slightly adherent. Outer lip with well developed internal denticles including ID, D1 to D6. D1 and D2 bifid. Outer lip not crenulate. Siphonal canal narrow, open, straight, up to 57% of apertural length. Pseudoumbilicus narrow.

Rouault (1850) described from Bos d’Arros (a locality very close to Gan), Murex ‌trigonus based on juvenile specimens. Unfortunately, Murex ‌trigonus ‌Rouault, 1850 is a junior homonym of Murex ‌trigonus ‌Gmelin, 1791 and Magne (1941), a homonymy as yet uncorrected. Therefore, we propose the new name Timbellus ‌magnei n. name herein. The juvenile specimens illustrated by Rouault (1850) are characterized by low intervarices varying from three to five (compare Fig. 12C, D and G). This character is present on the early whorls of the adults of T. ‌tripteroides ‌sensu Merle (1990: pl. 2, figs 13, 14 only). Thus, one can accept that these large T. ‌tripteroides ‌sensu Merle (1990) correspond to adult specimens of T. ‌magnei n. name. Timbellus ‌magnei n. name is easily separated from typical T. ‌tripteroides by having a multispiral protoconch (Fig. 12B), whilst typical tripteroides has a paucipiral protoconch (see Fig. 8D, E) and by having more and lower intervarices.

Cossmann & O’Gorman (1923) misinterpreted Murex ‌trigonus ‌sensu Rouault and illustrated a specimen typical of the genus Ponderia: P. ‌daguini ‌(Magne, 1941). In addition, Cossmann & O’Gorman (1923) described and figured an adult specimen of Timbellus that Cosmann named Murexcf. tricarinatus ‌Lamarck, 1803. Magne (1941) gave the name Murex(Pteropurpura)palensis to Murex cf. tricarinatus ‌Lamarck, 1803 (sensu Cossmann 1923) and demonstrated that the juvenile specimens illustrated by Rouault (1850) as Murex ‌trigonus were not juveniles of Murex (Pteropurpura) palensis. We discovered a young specimen from Gan differing from the juveniles of T. ‌magnei n. name. It displays one high intervarix between each varix (Fig. 12G), a character also present on the early teleoconch whorls of well preserved adults of T. palensis (Fig. 12H, I). Douvillé & O’Gorman (1929: pl. 29, fig. 1) figured a specimen from the early Eocene of Biron (near Orthez) and called it Murex ‌tripteroides. That specimen bears one high intervarix between each varix and possibly represents T. palensis.

Holotype. France• Paris Basin, Oise, Trosly-Breuil (Fond Couturier); Sables de Cuise, (biozone NP12); early Eocene (Ypresian, lower Cuisian); MNHN.F.A90544 (Six coll.), H: 14 mm (Figs 13; 26M).

Dedicated to Didier Six who collected this rare species.

Sables de Cuise, (biozone NP12), see Gély & Lorenz (1991: pl. 1), early Eocene (Ypresian, lower Cuisian).

France, Paris Basin, Oise, Trosly-Breuil (Fond Couturier).

Only known from the type locality.

Unknow protoconch. Teleoconch up to 14 mm in height, up to 7.6 mm in width, biconic in profile, composed of five whorls. Moderately high spire. Last whorl 74% of total length. Apical angle 74° including spines, 66° excluding spines. Spiral sculpture with fine, narrow, well developed primary cords. First whorl, no spiral sculpture. Second whorl, appearance of weak P1 and P2. Third whorl, P1 and P2 well marked, appearance of s1. Fourth whorl, appearance of threads on infrasutural ramp. Fifth whorl: P1 to P6, s1 to s3; primary and secondary cords almost same relief; ADP to ABP on siphonal canal. On first whorl, nine protovarices. On second whorl, appearance of varices with two intervarices intercalated. On third whorl: three varices and three intervarices intercalated. From fourth to fifth whorl, three varices and two or three intervarices intercalated. Intevarices low. Higher relief of the intervarices on P2. Varices sublamellose on early whorls, rather thick on last whorls. Appearance of short P1 spine on second whorl. On last whorl, short P1 spine oriented adapically; no other cord spines. Ovate aperture up to 34% of diameter and up to 68% of length of last whorl (including the siphonal canal). Columellar lip smooth, narrow, slightly erect anteriorly, forming flat curved inductura at base. Parietal lip adherent. Outer lip with well developed denticles including ID, D1 to D3 (D4-D5 probably not formed); all denticles simple. Siphonal canal open, slightly dorsally recurved, up to 43% of apertural length. Pseudoumbilicus narrow.

This shell represents the oldest known Timbellus from the Paris Basin, as no Timbellus has been recorded from the Early Cuisian (biozone NP12, Ypresian) despite intensive fieldwork. It is easily distinguishable from the middle Eocene Paris Basin Timbellus species: T. tripteroides ‌(Lamarck, 1822), T. ‌crenulatus ‌tricarinatus ‌(Lamarck, 1803), T. ‌micropterus ‌(Deshayes, 1835) and T. ‌capitaneus ‌Pacaud, Ledon & Goret, 2017. Juveniles of T. tripteroides and T. ‌capitaneus have weak spiral sculpture; Timbellus ‌crenulatus ‌tricarinatus lacks internal denticles, and juveniles of T. ‌micropterus have coarser spiral sculpture. Timbellussixi ‌n. sp. resembles juveniles of T. ‌magnei n. name from Bos d’Arros and Gan (Fig. 12C, D) in sharing a similar shape of its cords (fine, narrow, but well marked) and by having flat intervarices between the varices. However, T.sixi ‌n. sp. differs from T. ‌magnei n. name in having less foliated varices, having less numerous intervarices (usually 2-3 instead 4-5 in T. ‌magnei n. name), and by having MP and ABP on the siphonal canal.

In this species group, the last whorl is generally broader than in the species group of T. tripteroidesand the shoulder spine (P1) is more strongly developed. The spiral sculpture corresponds to fine primary cords, as in the T. tripteroides species group. Two species have an acute open shoulder spine, similar to that seen in the genus Pterochelus ‌Jousseaume, 1880: the Eocene species T.crenulatus ‌(Röding, 1798) and the Miocene species T.valdintortus ‌(Cossmann & Peyrot, 1924). In the T. ‌crenulatus species group the rounded aperture bears poorly developed denticles or lirae within the outer lip, except in the oldest known member (T. palensis[Magne, 1941]) from the early Eocene of Aquitaine Basin of Gan (Fig. 12G-I). However, according Cossmann & O’Gorman (1923), the short shape and the rounded aperture of shells of this species resemble more closely T. ‌crenulatus ‌tricarinatus ‌(Lamarck, 1803) than T. tripteroides. Probably, the presence of internal denticles in T. palensis represents a plesiomorphic character state and its loss in younger species of this group (e.g., T. ‌crenulatus ‌tricarinatus and T. ‌crenulatus crenulatus) represents a derived state. For this group, we describe a new species: T. ‌curvispina ‌n. sp. from the Lutetian of the Paris Basin. As for T. tripteroides, the type material of Murex ‌tricarinatus ‌Lamarck, 1803 is lost, as it is also missing in the collections of Geneva and Paris. In order to avoid future taxonomic mistakes, we designate a neotype herein (MNHN.F.A90547, Fig. 14B, C) from the original type locality of Grignon (precisely Falunière of Grignon; Calcaire à Orbitolites complanatus Formation, biozone NP15, see Gély & Lorenz [1991: pl. 1], middle Lutetian).

Holotype. France• Paris Basin, Oise, Parnes (Les Bôves); Falun de Parnes (biozone NP15); middle Eocene (middle Lutetian); MNHN.F.A90541 (ex B63594), H: 51.8 mm (Figs 14E, F; 26J).

Paratypes. France• 3 spm Paris Basin; Eure; Fontenay-en-Vexin (Bois du but); Falun de Parnes (biozone NP15); middle Eocene (Lutetian); MNHN.F.A73848 (Figs 14D; 26L), MNHN.F.A90542, MNHN.F.A90543 (Pons coll.) (Figs 14G, H, 26K)• 1 spm; idem; MNHN.F.A32514 (Pacaud coll.), figured inMerle et al. (2011: 440, pl. 98, fig. 7)•1 spm; Paris Basin, Yvelines, Villiers-Saint-Frédéric (Butte Saint-Léonard); Falun de Parnes (biozone NP15); Lutetian; MNHN.F.A32516 (Schtrock coll.), figured inMerle et al. (2011: 440 pl. 98, fig. 6)• 1 spm; Paris Basin, Yvelines, Parnes; Falun de Parnes (biozone NP15); Lutetian; MNHN.F.A90544 (ex Galerie de Zoologie coll.).

Name derived from Latin curv- (meaning curved) and spina (meaning spine) and refers to the P1 spine dorsally curved; gender female.

Falun de Parnes (biozone NP15), see Gély & Lorenz (1991: pl. 1), middle Eocene (middle Lutetian).

France, Paris Basin, Oise, Parnes (Les Bôves), see Fritel (1910: 221).

Calcaire grossier moyen, biozone NP15, Paris Basin (France), middle Eocene (middle Lutetian).

Protoconch not preserved. Teleoconch up to 51.8 mm in height, up to 30 mm in width, subfusiform in profile, composed of six whorls. Moderately high spire. Last whorl up to 85% of total length. Apical angle 63°. Spiral sculpture with marked primary cords on varices, but weaker on intervarices. First and second whorls eroded. From third: appearance of weak P1 and P2. Third word: no change. Fourth whorl: appearance of s1. Fifth whorl: occasional appearance of s2. Sixth whorl: P1 to P6, s1, s2, P6 not atrophied, s1 well developed; ADP developed, MP, ABP weak or missing on siphonal canal. Axial sculpture eroded on two early whorls. On second whorl, eight protovarices. Third whorl: appearance of varices; three varices and one intervarix between. From fourth to last whorl, no change. Higher relief of intervarices on P2. Varices sublamellose. Appearance of short straight P1 spine on fourth whorl. On last whorl, axial ornamentation forming very short spinelets, dorsally recurved on P1 to P6. P1 spine more strongly developed than other cord spines. Rounded aperture up to 34% of diameter and up to 74% 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 very weak denticles including D1 to D6. Small crenulations at edge of outer lip. Siphonal canal narrow, open, bent, slightly dorsally recurved, up to 81% of apertural length. Pseudoumbilicus narrow

This species can be included in the group of T. ‌crenulatus defined by Harasewych & Jensen (1979), Vokes (1992), and updated by Merle et al. (2011: 15). In this group the last whorl is generally wider than in species of the T. tripteroides group, the shoulder spine (P1) is more delineated and the internal denticles within the outer lip are weaker. The spiral sculpture corresponds to fine primary cords, as in the T. tripteroides group. Timbellus ‌curvispina ‌n. sp. was mistaken for the Lutetian species T. ‌crenulatus ‌tricarinatus ‌(Lamarck, 1803) but it differs by the construction of the P1 spine. This spine is straight, long and oriented abaperturally in T. ‌crenulatus ‌tricarinatus, whereas it is shorter, curved and dorsally oriented in T. ‌curvispina ‌n. sp. (see Fig. 14). Timbellus ‌curvispina ‌n. sp. also bears cord spines that are always more developed than in T. ‌crenulatus ‌tricarinatus. Moreover, the spiral cords of T. ‌curvispina ‌n. sp. are weak in the intervarical spaces, whereas those of T. ‌crenulatus ‌tricarinatus are well marked. Pacaud et al. (2017) recently designated the lectotype of Murex ‌tricarinoides ‌Deshayes, 1835 demonstrating that authors confused this species with another: T. ‌capitaneus ‌Pacaud, Ledon & Goret, 2017. In addition, they synonymised T. tricarinoides with T. ‌crenulatus ‌tricarinatus. However, T. tricarinoides differs from T. ‌crenulatus ‌tricarinatus in lacking true P1 spines and in displaying a rounded outer lip. T. ‌crenulatus crenulatus ‌(Röding, 1798) from the Bartonian is easily distinguishable from T. ‌curvispina ‌n. sp. by its acute and long P1 spine pointing abaperturally.

Members of this group are usually large sized and share a strong development of the internal denticles within the outer lip and strong primary cords; secondary cords are poorly developed. All species lack columellar denticles, except in the holotype of T.xenos ‌(Harasewych, 1982), which is presumably a pathological specimen (Vokes 1992: 18). In the middle Eocene from Europe, Timbellus ‌submicropterus ‌n. sp. (upper Lutetian) and the Bartonian T. ‌micropterus ‌(Deshayes, 1835) represent the oldest members of this species group. It becomes more diversified in the late Eocene/early Oligocene with T. ‌lamarckii ‌(Grateloup, 1845), T. ‌detritus ‌(von Koenen, 1890), T. ‌arenarius ‌(Steuer, 1912), T. ‌danapris ‌Pacaud, 2018 and T. ‌radulfiensis ‌n. sp. It contains three Miocene species: T. ‌perlongus ‌(Bellardi, 1872), T. ‌rosavendae ‌(Bellardi, 1872) and T. ‌pseudolamarcki ‌(Magne, 1941), and becomes extinct before the Pliocene.

Holotype. France• Normandy, Manche, Rauville-la-Place; Argiles à Corbules Formation; late Eocene (Priabonian); MNHN.F.A77754 (MNHN coll.), H: 13.5 mm (Figs 15A, B; 26N).

Paratypes.France• 1 spm; same as for the holotype; MNHN.F.A77755 (MNHN coll.) (Figs 15C; 26O)•3 spm; idem; MNHN.F.A77756 (MNHN coll.)•3 spm; idem; MNHN.F.A24118 (Vasseur coll.)•1 spm; idem; MNHN.F.A90559 (Vasseur coll.) (Figs 15D; 26P).

From the ancient name of the type locality Radulfi villa around 1083.

Argiles à Corbules Formation, late Eocene (Priabonian), stratigraphic data given by Cavelier (1979) and Chateauneuf (1980).

France, Normandy, Manche, Rauville-la-Place (Cauvin), Cavelier (1979).

Only know from the type locality.

Bulbous protoconch of one whorl. Teleoconch up to 20.5 mm in height, up to 7.5 mm in width, subfusiform in profile, composed of six whorls. Moderately high spire. Last whorl up to 74% of total length. Apical angle 61°. Spiral sculpture with moderately marked primary cords. First whorl: no cord. Second whorl: appearance of P2. Third whorl: appearance of P1 (weaker than P2) and s2. Fourth and fifth whorl: no change. Sixth whorl: P1 to P6, s1; P6 not atrophied; ADP, MP and ABP on siphonal canal. From first whorl to second whorl: ten protovarices. On third whorl: seven protovarices. Fourth whorl: appearance of varices; three varices and one intervarix intercalated. From fifth to sixth whorl: no change. Higher relief of intervarices on P2. Varices sublamellose. Appearance of short P1 spine on third whorl. Abapically spinelets from P1 to P6 and ADP. Ovate aperture up to 34% of diameter and up to 74% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, poorly erect anteriorly, forming curved inductura at base. Parietal lip slightly adherent. Outer lip with developed denticles including ID, D1 to D6. Outer lip not crenulate. Siphonal canal open, straight, up to 47% of aperture length. Pseudoumbilicus narrow.

This species can be compared to other Priabonian and Rupelian species from Northern Europe (Germany) and Eastern Europe (Ukraine) with which it shares a narrow shape. It differs from T detritus ‌(von Koenen, 1890), T. ‌arenarius ‌(Steuer, 1912) and T. ‌danapris ‌Pacaud (2018) in having finer spiral cords. The spiral sculpture resembles more that of T. ‌priabonicus ‌Pacaud, 2018 (nom. n. for Murex ‌trialatus ‌von Koenen, 1889, non G. B. Sowerby (II), 1834), but T. ‌priabonicus bears strongly developed P1 and P2 spines, whereas only P1 spine is developed in T. ‌radulfiensis ‌n. sp.

Holotype. France• Normandy, Manche, ­Fresville; Calcaire de Fresville; middle Eocene (uppermost Lutetian); MNHN.F.A24111 (MNHN coll.), H: 23.6 mm (Figs 16A, B; 27A).

Paratypes. France• 1 spm; same as for the holotype; UCBL-EM 33438 (Bourdot coll., figured specimen of Cossmann & Pissarro 1901) (Figs 16E, F; 27C)•1 spm; idem; MNHN.F.A90556 (MNHN coll.) (Fig. 16K)•1 spm; idem; MNHN.F.B63700 (de Morgan coll.)•1 spm; Normandy, Manche, Gourbesville; Calcaire de Fresville; upper Lutetian; MNHN.F.A24126 (Munier-Chalmas coll.), figured specimen of Merle et al. (2011: 135, pl. 102, fig. 4a, b) (Figs 16C, D; 27B).

Additional material (material referring to Timbellus cf. ‌submicropterus ‌n. sp.)

France• 3 spm; Loire-Atlantique, Loire Basin, Saffré (Bois Gouët); middle Eocene (lower Bartonian); MNHN.F.A29660 (Pacaud coll.)•2 spm; Saint-Aignan-Grand-Lieu (Pierre-Aiguë), Loire-Atlantique; idem; MNHN.F.A29662, (Brébion coll.), MNHN.F.A90557 (Brébion coll. (Figs 16G, H; 27D).

Name formed by the prefix sub- and micropterus for this species older than Bartonian Timbellus ‌micropterus (­Deshayes, 1835).

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

France, Normandy, Manche, Fresville, see APGN (2024).

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

Bulbous protoconch of one whorl (Fig. 16K). Teleoconch up to 23.6 mm in height, up to 11.6 mm in width, biconic in profile, composed of six whorls. Moderately high spire. Last whorl up to 70% of total length. Apical angle 59°. Spiral sculpture with coarse primary cords. First to third whorl: no cord. Fourth whorl: appearance of weak P1, s1 and P2. Fifth whorl: P1, s2 and P2 coarser. Sixth whorl: P1 to P6, s1; P6 not atrophied on convex part of whorl; ADP, MP and ABP on siphonal canal. From first whorl to second whorl: 6-7 protovarices. On third whorl: appearance of varices; three varices and one intervarix intercalated. From third to sixth whorl: no change. Higher relief of intervarices on P2. Varices rather thick. Appearance of short P1 spine on third whorl. Ovate aperture up to 27% of diameter and up to 71% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, poorly erect anteriorly, forming curved inductura at base. Parietal lip slightly adherent. Outer lip with developed denticles including ID, D1 to D6. D6 in some specimens. Outer lip not crenulate. Siphonal canal open, straight, up to 40% of apertural length. Pseudoumbilicus narrow.

A specimen of this species (Figs 16E, F; 27C) was identified as Murex(Pterymurex)tricarinatus ‌Lamarck, 1803 (synonym of Timbellus ‌crenulatus ‌tricarinatus) by Cossmann & Pissarro (1901). Timbellus ‌crenulatus ‌tricarinatus differs greatly by apertural (with weak internal denticles within the outer lip), sculptural and shape characters, and belongs to another group species (see above Fig. 14). In fact, this specimen illustrated by Cossmann & Pissarro (1901) more closely resembles T. ‌micropterus from the Bartonian Paris Basin. Merle et al. (2011) attributed another Norman specimen (pl. 102, fig. 4a, b) to T. ‌micropterus (Figs 16C, D; 27B). It is true that these two specimens are superficially similar to T. ‌micropterus, but further comparison reveals some differences, such as only one intervarice (instead two in typical T. ‌micropterus) and finer secondary cords on the last whorl. The bivariate plot of total length against width (Fig. 17) does not suggest strong differences Normandy and Paris Basin populations. Finally, it is tempting to conclude that the observed teleoconch differences in T. ‌submicropterus ‌n. sp. fall within the intraspecific variation of T. ‌micropterus. However, T. ‌submicropterus ‌n. sp. has a paucispiral protoconch, whereas T. ‌micropterus has a multispiral (Fig. 16K, L), suggesting two separate species with different modes of larval development. In the Bartonian from Loire-Atlantique, there are specimens showing coarse spiral cords like those of typical T. ‌submicropterus ‌n. sp., but they bear a more pronounced shoulder (Fig. 16G, H). Their protoconch is not preserved. We therefore attribute these specimens to T. ‌submicropterus ‌n. sp. with hesitation.

Murex ‌pinnatus ‌Swainson, 1822, by monotypy. Recent: Indo-Pacific.

As seen above, fossil and Recent members of the genus Timbellus ‌de Gregorio, 1885 were usually attributed to Pterynotus ‌Swainson, 1833 (e.g., Cossmann 1889; Cossmann & Pissarro 1911; Glibert 1963; Vokes 1970) in the subfamily Muricinae. Based on molecular data, Barco et al. (2010, 2012) studied the phylogenetic relationships of Pterynotus represented byP. ‌elongatus ‌(Lightfoot, 1786) and P. ‌fulgens ‌Houart, 1988 and Pterymarchia ‌Houart, 1995 represented by P. ‌martinetana ‌(Röding, 1798). These authors demonstrated firstly, that Pterynotus and Pterymarchia are not phylogenetically close to P. ‌fulgens ‌Houart, 1988 and secondly, none of these taxa are included within the clade of the Muricinae. According to Merle et al. (2011), Pterynotus ‌fulgens is closely related to the type species of Timbellus and is now attributed to that genus. Thus, Merle et al. (2011) and Houart (2018) rejected Pterynotus and Pterymarchia from the Muricinae. Russini et al. (2023: 866) place Pterynotus in incertae sedis and consider that it likely represented an independent lineage, worthy of subfamilial rank.

Murex ‌tripterus ‌Born, 1778, by original designation. Recent: Indo-Pacific.

According to Houart (1995), Pterymarchia can be distinguished from Pterynotus (s.s.) by having more numerous axial varices on the early teleoconch whorls (9-11 vs 6-8), later appearance in ontogeny of the trivaricate sculpture, when present, sometimes more varices on the last whorls (3-5 vs 3), and more strongly developed denticles. Houart (1995, 2018) proposed full generic status for Pterymarchia. However, we prefer to maintain it at subgeneric rank because: 1) based on molecular data, Pterynotus and Pterymarchia are very close (Barco et al. 2010); and 2) based on shell characters, Pterymarchia shares with Pterynotus a scabrous surface appearing early in ontogeny, generally fine to moderately fine primary cords (from IP to ABP), and a series of internal denticles from ID to D6. Furthermore, several other characters are occasionally shared by both genera, such as the trivaricate sculpture, columellar tubercles [e.g., P. (s.s.) patagiatus (Hedley, 1912), P. (s.s.) praepatagiatus Vokes, 1992 and P. (s.s.) granuliferus (Grateloup, 1833)], and even developed denticles in some P. (s.s.) albobruneus Bertsch & D’Attilio, 1980.

Holotype. France• Aquitaine Basin, Landes, Gaas (Espibos); Gaas Formation (biozone NP23); early Oligocene (Rupelian); MNHN.F.A32492 (Merle coll.), figured specimen in Merle et al. (2011: pl. 90, fig. 3), H: 27 mm (Figs 18A, B; 27E).

Paratypes.France• 6 spm; same as for the holotype; MNHN.F.A77749 (Merle coll.)•1 spm; idem; MNHN.F.A90554 (Merle coll.) (Figs 18C, D; 27F)•1 spm; idem; MNHN.F.A86913 (Lozouet coll.), figured in Lozouet (2023: text-fig. 10a, b).

France • 1 spm; Aquitaine Basin, Landes, Gaas (La Gouarde); same as for the holotype; Aucoin coll., figured inMerle et al. (2011: pl. 90, fig. 4a, b).

From the type locality, Gaas (Landes), France.

Gaas Formation (biozone NP23), see Sztrákos & Steurbaut (2017: 752), early Oligocene (Rupelian).

France, Aquitaine Basin, Landes, Gaas (Espibos), see Sztrákos & Steurbaut (2017: 752).

Aquitaine Basin (France), early Oligocene (Rupelian).

Paucispiral protoconch (fide Lozouet 2023: text-fig. 10e). Teleoconch up to 27 mm in height, up to 12 mm in width, biconic in profile, composed of six whorls. Moderately high spire, first two whorls subcarinate; next whorls rounded. Last whorl up to 75% of total length. Apical angle 56°. Spiral sculpture with marked primary cords. First whorl: appearance of primary cords P1 and P2; second whorl: P1 and P2; third whorl: appearance of s1. Fourth whorl: appearance of abis, adis and s2. Fifth whorl: no change or appearance of threads. Sixth whorl: IP, adis and abis, P1 to P6, s1 to s6; s2 and s3 well developed; P6 on base on convex part of whorl; ADP, MP and ABP slightly more developed than ads and ms. Surface of shell with numerous threads appearing on fourth whorl. Axial sculpture with thick varices. On first whorl: nine lamellose varices. From the second to fifth whorl: 9-10 thick varices. On sixth: appearance of three or four varices and one intervarices intercalated. Varices and intervarices thick. From the third to last whorl, numerous, fine microlamellae covering entire surface forming scabrous microsculpture. Ovate aperture up to 25% of diameter and up to 66% of length of last whorl (including siphonal canal). Columellar lip, adherent posteriorly, becoming progressively more erect anteriorly. One or two small columellar tubercles. Parietal lip adherent. Outer lip bearing strong internal denticles: ID, D2 to D6 present, D1 missing. Pseudoumbilicus moderately large. Siphonal canal open, up to 48% of apertural length, slightly dorsally recurved.

Merleet al. (2011: pl. 90, figs 3-4) attributed with hesitation two specimens from Gaas to Murex ‌subquadrifrons ‌d’Orbigny, 1852, originally described as Murex ‌quadrifrons by Grateloup (1845) from this same Rupelian locality. The Grateloup’s collection is housed in the University of Talence, but the type material of M. ‌quadrifrons was not found and is likely lost (personal observation of the first author). Therefore, only the drawing of Grateloup (1845: pl. 3 [pl. 31], fig. 10, refigured in Merle et al. 2011: pl. 90, fig. 2) can be used to identify the species. According to Merle et al. (2011), the drawing shows a shell with a textiliose surface and a biconic shape, as in the Gaas material at hand. Moreover, to our knowledge, no other muricid from Gaas except P. (Pterymarchia) is comparable to the specimen illustrated by Grateloup (1845). However, the shell illustrated by Grateloup (1845) lacks internal denticles within the outer lip, lacks columellar denticles, shows more numerous varices (six on the ventral face vs four in the specimens at hand), has a narrower siphonal canal, and is more likely to represent a specimen of Favartia. These numerous differences between Grateloup’s (1845) figure and the studied material suggest that it obviously corresponds to a species other than M. ‌subquadrifrons. Murex ‌subquadrifrons is therefore not recognizable using the figure of Grateloup, lacks type material and it should be considered a nomen dubium.

Eocene Pterynotus(Pterymarchia), such as P.(P.)antiquus ‌(Merle 1990), P.(P.)denudatus ‌(Deshayes, 1835) and P.(P.)defensus ‌(Fuchs, 1870) strongly differ from the P.(P.)gaasensis ‌n. sp. in having trivaricate sculpture. Pterynotus(P.)fascistria ‌(von Koenen, 1889) from the Priabonian of North Germany is more similar shape, as it lacks trivaricate sculpture and because their primary cords are poorly distinguishable from the secondary cords on the last whorls. Pterynotus(P.)fascistria is distinguished from P.(P.)gaasensis ‌n. sp. in having a lower spire and by lacking intervarices, D1 and ID denticles. Pterynotus(P.)gaasensis ‌n. sp. can be compared to two other Neogene species: P.(P.)pelouatensis ‌n. sp. from the Burdigalian of Aquitaine Basin (Figs 18E-H; 27G, H) and P.(P.)sandbergeri ‌(Hörnes, 1856) from the Langhian of the Paratethys. Both species differs from P.(P.)gaasensis ‌n. sp. in having more angular whorls with a shoulder delimited by a subspiny P1 and by other primary cords more distinguishable from the secondary cords on the last whorl. Lozouet (2023) followed Merle et al. (2011) and adopted the name Pterynotuscf. subquadrifrons for the Rupelian material from Gaas, but he did not include Chattian and Aquitanian specimens figured in Merle et al. (2011: pl. 90, figs 5, 6), because Rupelian shells have a paucispiral protoconch, whereas the Chattian/Aquitanian species has a multispiral protoconch, and is therefore a separate species. Thus, we consider for instance that P.(P.)gaasensis ‌n. sp. to be present only in the Rupelian.

Holotype. France• Aquitaine Basin, Gironde, Saucats (Peloua); Faluns de Saucats; Early Miocene (lower Burdigalian); MNHN.F.J09945 (Neuville coll.), figured specimen in Merle et al. (2011: 424, pl. 90, fig. 7), H: 21 mm (Figs 18E, F; 27G).

Paratype.France• 1 spm; same as for the holotype; MNHN.F.J17632 (Neuville coll.) (Figs 18G, H; 27H).

From the type locality, Peloua, Gironde, France.

Faluns de Saucats, Early Miocene (lower Burdigalian), see Parize et al. (2008: 396).

France, Aquitaine Basin, Gironde, Saucats (­Peloua), see Parize et al. (2008: fig. 1).

Only known from the type locality.

Two smooth protoconch whorls preserved delimited by sinusigera scar (protoconch tip not preserved) (see Merle et al. 2011: 122, text-fig. 46E). Teleoconch up to 21 mm in height, up to 12.5 mm in width, biconic in profile, composed of five whorls. Moderately high spire with subcarinate whorls. Last whorl up to 73% of total length. Apical angle 64°. Spiral sculpture with marked primary cords. First whorl: appearance of P1 and P2; second whorl: appearance of IP; third whorl: appearance of abis, adis and s2. Fourth whorl: no change or appearance of threads. Fifth whorl: IP, adis and abis, P1 to P6, s1 to s6; s3 and s4 more developed than other secondary cords; P6 slightly atrophied, on base of convex part of whorl; ADP, MP, ads and ms; ABP missing. Surface of shell with numerous threads appearing on fourth whorl. Axial sculpture with thick varices. On first whorl: 10-11 lamellose varices; on second whorl: 10-12 varices; on third whorl: 11-12 varices; on fourth whorl: 10-12 varices; on fifth whorl 8-10 varices. From second whorl to fifth whorl: thick varices. No evidence of intervarices on last whorl. From third to last whorl numerous, fine microlamellae forming scabrous microsculpture covering entire surface. Ovate aperture up to 31% of diameter and up to 72% of length of last whorl (including siphonal canal). Columellar lip adherent posteriorly, becoming progressively more erect anteriorly. One or two weak columellar tubercles. Parietal lip adherent. Outer lip with strong internal denticles: ID, D2 to D6 present, D1 missing; D4 split, D6 poorly developed. Pseudoumbilicus moderately large. Siphonal canal open, up to 43% of aperture length, slightly dorsally recurved.

The European Neogene contains two species of Pterynotus(Pterymarchia): P. (P.)pelouatensis ‌n. sp. and P.(P.)sandbergeri ‌(Hörnes, 1856) ‌n. comb. from the Langhian of Paratethys (Austria, Hungary, Romania, Poland) and the Serravallian of Turkey. The type material of P.(P.)sandbergeri is housed in Naturhistorisches Museum of Wien and contains the figured syntype of the species (Hörnes 1856: pl. 51, fig. 5a, b, NHMW 1855/0045/0435). Here, we designate this specimen as the lectotype (Fig. 18I, J). Pterynotus(P.)sandbergeri was often attributed to Murexsul ‌Iredale, 1915 (Landau et al. 2013; Kovács et al. 2018; Kovács 2019). Although they share several similarities (biconic shells, internal denticles, columellar denticles), members of Murexsul have a more rounded aperture, their internal denticles are close together, ID is weaker and their siphonal canal is narrower. The spires of P.(P.)pelouatensis ‌n. sp. and the lectotype of P.(P.)sandbergeri are very similar, but their last whorls are very different. The lectotype of P.(P.)sandbergeri has a larger aperture, a surface covered by numerous threads and atrophy of P2, P5 and P6; P4 is more developed and D6 is missing. The differences between the primary cords are great, but in other specimens attributed from P.(P.)sandbergeri (Fig. 18K, L), the atrophy of P2, P5 and P6 is less strong than in the lectotype from Austria (see Landau et al. 2013; Kovács et al. 2018; Kovács 2019). Compared to extant P. (Pterymarchia), P.(P.)pelouatensis ‌n. sp. and P.(P.)sandbergeri share greatest morphological similarities with P.(P.)martinetana ‌(Röding, 1798) and particularly with specimens from the Red Sea (Merle et al. 2011: pl. 92, figs 11, 12), but P.(P.)sandbergeri differs by having develop cord spines on P1, P2, P4, P5, ADP and MP.

Murex ‌textiliosus ‌Lamarck, 1803 by original designation. Middle Eocene: Paris basin, France and Hampshire Basin, England.

Textiliomurex shares similarities with Pterynotus (s.s.) and Pterynotus (Pterymarchia) such as a scabrous surface, columellar denticles and a series of internal denticles including ID, D1 to D6. Textiliomurex does not display trivaricate axial sculpture and lacks intervarices, usually found in Pterynotus (s.s.). The number of varices on the last whorls of Textiliomurex is higher (9-11) than in Pterymarchia (3-5, see Houart 1995). As with Pterynotus, we cannot give a subfamilial placement to this genus. Merle et al. (2011: 127) included six species in the genus. Recently, Lozouet (2023) added (with some doubt) one species from the Chattian of the Aquitaine Basin (Textiliomurex?problematicus ‌Lozouet, 2023), and we add a new species from the Lutetian of the Paris Basin herein.

Holotype. France• Paris Basin, Oise, Chaumont-en-Vexin (Carrière Darcy); Calcaire Grossier inférieur (biozone NP14); middle Eocene (lower Lutetian); MNHN.F.A81144 (Pacaud coll.), H: 28 mm (Figs 19A, B; 27I).

Paratype. France• 1 spm; same as for the holotype; MNHN.F.A81145 (Pons coll.) (Figs 19C, D; 27J).

Name from the type locality Chaumont-en-Vexin. This place name comes from the Celtic words Chod “wood” and Mon “montain”.

Calcaire Grossier inférieur (biozone NP14), see Gély & Lorenz (1991: pl. 1), middle Eocene (lower Lutetian).

France, Paris Basin, Oise, Chaumont-en-Vexin (Carrière Darcy), see Ott d’Estevou et al. (2014).

Only known from the type locality.

Protoconch not preserved. Fusiform teleoconch up to 28 mm in height, up to 21 mm in width. High spire, up to seven rounded whorls. Last whorl (seventh whorl) up to 65% of total length. Apical angle 55°. Spiral sculpture with fine, narrow, well marked primary and secondary cords. First to third whorl not preserved. Fourth whorl: IP, s1 and P2. Fifth whorl: appearance of the secondary cord adis (sutural ramp). Sixth whorl: addition of tertiary cords on sutural ramp and convex part of whorl. Seventh and last whorl: IP, adis and abis (sutural ramp), P1 to P6 (convex part of the whorl); P6 placed on base on convex part of whorl; ADP, MP, ABP (siphonal canal), MP, ABP poorly distinct from secondary cords; primary and secondary cords of same thickness. Surface of shell with numerous threads. Axial sculpture with nine varices on fourth whorl, seven on fifth whorls, six on sixth whorls and five on seventh and last whorl. Nodulose varices. No intervarices. No varical spine. Numerous, fine microlamellae covering entire surface forming scabrous microsculpture. Ovate aperture up to 36% of diameter and up to 70% of length of last whorl (including siphonal canal). Columellar lip adherent, smooth. Parietal lip adherent. Short anal sulcus. No shoulder channel. Outer lip with subobsolete internal denticles. Outer lip finely crenulated. Pseudoumbilicus wide. Siphonal canal open, up to 38% of apertural length, slightly dorsally recurved.

Textiliomurex ‌chodmonensis ‌n. sp. can be compared to two other species from the middle Lutetian: T. textiliosus ‌(Lamarck, 1803) and T. ‌bernayi ‌(Deshayes, 1865). Textiliomurex ‌textiliosus differs in having a shorter spire, a more inflated last whorl, a wider aperture, and a longer siphonal canal bearing a narrower pseudoumbilicus. Textiliomurex ‌bernayi has angular teleoconch whorls with a marked P1 shoulder, bears well developed internal denticles within the outer lip and cannot be confused with T. ‌chodmonensis ‌n. sp. (see Merle et al. 2011: 126, text. fig. 47 and pl. 94 about Textiliomurex).

Typhis ‌zealandica ‌Hutton, 1873 by original designation. Pleistocene to Recent: New Zealand.

Subfamilial placement of the genus Ponderia ‌Houart, 1986 has varied in the literature. D’Attilio & Hertz (1988) placed it in the subfamily Tripterotyphinae ‌D’Attilio & Hertz, 1988, whereas Vokes (1996) preferred placement it in the subfamily Muricinae. More recently, Merle et al. (2011) included Ponderia in a basal muricid group (without subfamilial assignment) and excluded it from the typical Muricinae. Houart (2018) included five genera in the Tripterotyphinae: Cinclidotyphis ‌DuShane, 1969, Prototyphis ‌Ponder, 1972, Pterotyphis ‌Jousseaume, 1880, Tripterotyphis ‌Pilsbry & Lowe, 1932 and Ponderia. All these genera share trivarical morphology, but the construction of the anal tube in Cynclidotyphis and Pterotyphis differs from that of Prototyphis, Ponderia and Tripterotyphis. In Cynclidotyphis and Pterotyphis, the anal tube is built just below the suture probably not with P1, whereas in Prototyphis, Ponderia and Tripterotyphis it is clearly built with P1. Tripterotyphis also differs from Ponderia and Prototyphis in having more strongly developed spiral sculpture. Thus, this group of five genera is likely to be polyphyletic. In their molecular phylogenetic work, Russini et al. (2023) confirm this hypothesis showing that: 1) Tripterotyphis belongs to a clade containing Muricopsis; and 2) Ponderia may represent a plesiomorphic lineage at the base of the pagoduline-haustrine diversification. Thus, those authors (p. 866) placed Ponderia in incertae sedis, and considered it likely to represent an independent lineage worthy of subfamilial rank.

Holotype. France• Normandy, Manche, Fresville; Calcaire de Fresville; middle Eocene (uppermost Lutetian); MNHN.F.A90548 (Remy coll.), H: 26 mm (Figs 19E, F; 27K).

Paratypes. France• 5 spm; same as for the holotype; MNHN.F.A90549 (Remy coll.)•1 spm; idem; MNHN.F.A90555 (Remy coll.) (Fig. 19G, H; 27L)•3 spm, MNHN.F.A90550 (MNHN coll.)•1 spm; Normandy, Manche, Fresville (coteau de Vauville); idem; MNHN.F.A27328 (MNHN coll.)•2 spm; idem; MNHN.F.B63590 (de Morgan coll.).

Dedicated to Nicolas Remy who collected the material of this species.

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

France, Normandy, Manche, Fresville, see APGN (2024).

Only known from the type locality.

Bulbous, paucispiral protoconch of 1.5 whorls. Slender, fusiform teleoconch up to 21 mm in height, up to 9 mm in width. High spire, up to six subcarinate whorls. Last whorl (sixth whorl) up to 69% of total length. Apical angle 35° excluding P1 spine and up to 55° including P1 spine. Spiral sculpture with strongly marked primary cords. First and second whorl: no primary cord. Third whorl: appearance of weak P2. Fourth whorl: P2 strongly marked and appearance of weak P1. Fifth whorl: strongly marked P1 and P2. Sixth and last whorl: P1 to P3 well marked, P4 weak (convex part of whorl); secondary cords occasionally s1 and s2. Axial sculpture with 8-9 lamellose protovarices per whorl from first to fourth whorl to four lamellose protovarices. From fifth whorl to sixth whorl, three major varices and two or three nodulose intervarices intercalated. Varices with three major cord spines (P1 to P3). P1 spine appearing at beginning of first whorl and forming open tube at end of first whorl. From second to last whorl P1 spine well developed, open. Intervarical nodules spirally aligned and present on P1 and P2. Ovate aperture up 32% of diameter (including wings) and up to 76% of length of last whorl (including siphonal canal). Columellar lip smooth, poorly erect. Parietal lip adherent. Shoulder channel of P1 open. Outer lip lacking denticle. Outer lip not crenulate. Pseudoumbilicus narrow. Siphonal canal open, up to 55% of apertural length.

Amongst European Eocene species, Ponderia ‌remyi ‌n. sp. can be compared to P. ‌daguini ‌(Magne, 1941) from the Early Eocene of Gan (Aquitaine and Paris Basins) and to P. ‌bispinosa ‌(Sowerby, 1823) from the Middle Eocene of the Paris Basin and Hampshire (England) and from the Late Eocene of Lattorf Basin (Germany). Ponderia ‌daguini is easily distinguished from P. ‌remyi ‌n. sp. in having a broader shape, more numerous cords (with P5, P6 and ADP) and nodulose cords. Ponderia ‌bispinosa usually bears more foliaceus varices. The cord spines of P. ‌bispinosa varies from two (P1 and P3) to five (P1 to P5). The morphotypes with five cords spines are common in the Lutetian and in the Bartonian of the Paris Basin, where those with two cord spines are common in the Bartonian of Hampshire and in the Priabonian of Lattorf (von Koenen 1889; Merle et al. 2011). The varices of P. ‌bispinosa are more winged than those of P. ‌remyi ‌n. sp. In addition, P. ‌bispinosa clearly differs in having a multispiral protoconch and a microsculpture formed of small elongated granules (see Merle 1990).

Flexopteron ‌philippinense ‌Shuto, 1969, by original designation. Late Miocene: Philippines.

The genus Flexopteron ‌Shuto, 1969 was previously included in the subfamily Muricinae (e.g., Vokes 1992, 1996; inter alia). According to Merle et al. (2011), it displays plesiomorphic shells, differing greatly from the derived shells found in typical Muricinae that are close to the genus Murex ‌Linnaeus, 1758. Merle et al. (2011: 54) defined an informal group called Basal muricid group including Flexopteron, Paziella, Poirieria, Timbellus, and several other genera. In a molecular phylogenetic work, Houart et al. (2021) found a clade including Flexopteron and Timbellus. This result could suggest close relationship between Flexopteron and Timbellus and confirms that they are not phylogenetically related to the derived clade of typical Muricinae. However, this relationship is not retrieved in the latest molecular phylogenetics of Russini et al. (2023), and those authors do not give subfamilial attribution to Flexopteron (incertae sedis position). Flexopteron has been considered in the past as a subgenus of Poirieria (e.g., Vokes 1992) or Paziella (e.g., Merle et al. 2011). As there is no evidence of a close phylogenetic relationship between these two genera based on molecular analyses, we give Flexopteron full generic status.

Holotype. France• Normandy, Manche, Rauville-la-Place (Cauvin); Argiles à Corbules Formation; late Eocene (Priabonian); MNHN.F.A77760 (MNHN coll.), H: 9.2 mm (fragment of last whorl) (Figs 20C, D; 27N).

Paratypes. France• 1 spm (juvenile); same as for the holotype; MNHN.F.A77761 (MNHN coll.), juvenile (Figs 20A, B; 27M)•10 spm (juveniles); idem; MNHN.F.A90551 (MNHN coll.).

From the Latin adjective Constantinus in reference to the type region of the species (pagus constantinus of the Gallo-Roman), the Cotentin (Manche).

Argiles à Corbules Formation, late Eocene (Priabonian), stratigraphic data given by Cavelier (1979) and Chateauneuf (1980).

France, Normandy, Manche, Rauville-la-Place (Cauvin), Cavelier (1979).

Normandy (France), late Eocene (Priabonian).

Smooth, paucispiral protoconch of 1.5 whorls. Teleoconch probably biconic (spire broken in most adult specimen), last whorl (holotype) up to 9 mm in height, up to 7 mm in width. Moderately high spire in juveniles. Rounded to subcarinate whorls. First whorl to fourth whorl: no cord. Fifth whorl: weak P1 to P5. P1 spine poorly developed. Axial sculpture with five sublamellose varices on first whorl, seven on second whorl, eight on third and fourth whorls. Ten varices on holotype. Ovate aperture, siphonal canal short. Columellar lip smooth, adherent posteriorly. No anal sulcus, no parietal callus. Outer lip with denticles D1, D2 and D3. Shoulder sinus of P1 spine shallow. Pseudo-umbilicus narrow. Siphonal canal short, open.

Although the material is composed of juvenile specimens (paratypes) and a last whorl of an adult specimen (holotype), it differs clearly from all Bartonian and Priabonian species from Northern Europe. The early whorls of Flexopteron ‌albionis ‌(Wrigley, 1930) from Barton beds, F.subplicatilis ‌(Wrigley, 1930) from the Bartonian of Southampton Dock and F.spinulosum ‌(Deshayes, 1835) from the Bartonian of the Paris Basin differ from those of F. ‌constantinense ‌n. sp. by having a more ornamented spiral sculpture with well developed P1 to P3. Flexopteronelatior ‌(von Koenen, 1889), found in the Bartonian of the Paris Basin and in the Priabonian of Lattorf (Germany) is another species bearing strong spiral ornamentation.

Holotype. France• Paris Basin, Oise, Liancourt-Saint-Pierre; Sables d’Hérouval (biozone NP12); early Eocene (Ypresian); MNHN.F.J02578 (Cossmann coll.), figured specimen in Cossmann & Pissarro (1911: pl. 39, fig. 169-17), H: 7.7 mm (Figs 20E, F; 28A).

Paratypes. France• 1 spm; same as for the holotype; MNHN.F.A77758 (Merle coll.) (Figs 20G, H; 28B)•17 spm; idem; MNHN.F.A77759 (Merle coll.)•9 spm; idem; MNHN.F.J17633 (Cossmann coll.);•1 spm; Paris Basin, Oise, Hérouval; Sables d’Hérouval (biozone NP12); early Eocene (Ypresian); MNHN.F.B63662 (Morlet coll.)•29 spm; idem; MNHN.F.A90913 (Pacaud coll.).

From the Latin adjective liancurtense in reference to the type locality, Liancourt-Saint-Pierre (Oise).

Sables d’Hérouval (biozone NP12), see Gély & Lorenz (1991: pl. 1), early Eocene (Ypresian).

France, Paris Basin, Oise, Liancourt-Saint-Pierre, see Fritel (1910: 190).

Paris Basin (France), early Eocene (Ypresian, Hérouval Formation only).

Smooth, conical multispiral protoconch of around three whorls. Sinusigeral scar well marked. Biconic teleoconch up to 7 mm in height, up to 4 mm in width (including cord spines). Moderately high spire, up to four subcarinate whorls. Last whorl (fourth whorl) up to 68% of total length. Apical angle 54° including P1 spine. Spiral sculpture with marked primary cords. First whorl: no cord; second whorl: appearance of P2 weak; third whorl: development of P1 spine and appearance of P3 above suture line. Fourth and last whorl: no primary cord on sutural ramp, P1 to P5, s1 and s2 moderately marked, P6 and ADP weak on siphonal canal. Axial sculpture with 11 lamellose varices on first whorl, 13 on second, and 14 on third and fourth. Varices axially aligned in a zig-zag fashion. Ovate aperture up to 36% of diameter and up to 77% of length of the last whorl (including siphonal canal). Columellar lip slightly erect anteriorly, adherent posteriorly, usually smooth or occasionnally with one tubercle. No anal sulcus, no parietal callus. Outer lip with small denticles. D1 to D5 simple, D1 more developed than other denticles. Outer lip not crenulate. Shoulder sinus of P1 spine widely open, shallow. Pseudoumbilicus narrow. Siphonal canal open, up to 36% of the apertural length and slightly dorsally recurved.

Melleville (1843) described Murex ‌foliaceous for a small Ypresian Flexopteron displaying axially zizgaging lamellae and a short P1 spine. Later, Cossmann (1889) described Murex ‌dyscritus which bears less marked primary cords than Murex ‌foliaceous. However, numerous specimens housed in the MNHN show many transitional forms suggesting these differences represent intraspecific variation. Thus, the older binome Murex ‌foliaceous takes nomenclatural priority. Moreover, Cossmann & Pissarro (1911) illustrated an atypic specimen from the Ypresian of Liancourt-Saint-Pierre (Hérouval Formation), squatter than usual for F. ‌foliaceum, with a bent siphonal canal, poorly developed P1 spines, more marked primary cords and occasionally a tubercle on the columellar lip. This specimen from Liancourt is not a pathological specimen and all specimens from this locality and Hérouval display similar morphology. They differ from F. ‌foliaceum (also found at Liancourt-Saint-Pierre) by having spiral cords P1 and P2 appearing earlier in ontogeny. Merle (1999, unpublished thesis) illustrated several other specimens and the ontogenetic changes in their spiral sculpture. He considered this population as a different species and named it Vauxyrytis liancurtensis. However, Merle (1999) is an unpublished thesis and the name is a nomen nudum made available herein. Placement of this species in the genus Vauxyrytis ‌Merle, 1999 (nomen nudum), an unpublished genus corresponding to Beyregrex ‌Merle ‌n. gen., (see below), was based on the shape of the varices, which are not spiny, and the occasional presence of a columellar tubercle. Nevertheless, we found some specimens of F. ‌liancurtense ‌n. sp. with P1 spines and several species of Flexopteron can have a columellar tubercle [e.g., F. ‌ogormani ‌(Cossmann, 1923)]. In addition, F. ‌liancurtense ‌n. sp. clearly resembles a Tortonian species from NW France: F. ‌gallicum ‌(Landau, Merle, Ceulemans & Van Dingenen, 2019), see Landau et al. (2019: pl. 11). Thus, we attribute this species to Flexopteron.

Murex ‌calcitrapoides ‌Lamarck, 1822 by original designation (synonym of Murex ‌calcitrapa ‌Lamarck, 1803). Middle Eocene (Lutetian and Bartonian): Paris and Loire basins, France.

Members of this genus display a biconic shell with a moderately high spire, spiny varices on P1, internal denticles (D1 to D6), a broad and open siphonal canal, a sinus of P1, around 8-10 varices, and complex spiral sculpture including the primary cords P1 to P6 (convex part of the whorl) and ADP-MP (siphonal canal). In addition, their shape is similar to that of Poirieria ‌Jousseaume, 1880 or Flexopteron (see Merle et al. 2011). For these reasons, Merle et al. (2011) placed Crassimurex (s. s.) and Crassimurex(Eopaziella) ‌Gürs, 2001 in an informal group named basal muricids. It is still impossible to place this genus within a subfamily using shell characters. Obervations on Crassimurex (s. s.) calcitrapa ‌Lamarck, 1803 at La Ferme de l’Orme (Lutetian, Paris Basin) suggest this species was a drilling predator (Merle et al. 2024), but this ability occurs in different subfamilies (e.g., Muricinae and Ocenebrinae) and does not help in subfamilial attribution.

Crassimurex(Pliocrassimurex)hirtus ‌n. sp., Early Pliocene (Zanclean): Spain.

Name formed by the beginning of Pliocene (Plio-) and Crassimurex.

Type species only.

Biconic, mid-sized teleoconch. Well marked primary cords. On spire P1 to P2 (convex part of whorl). On last whorl IP (infrasutural ramp), P1 to P5 (convex part of whorl), P6, ADP, MP and ABP (siphonal canal). P2 delimiting shell periphery. Varices lamellose on infrasutural ramp, spiny from shoulder (P1) to siphonal canal. Varices aligned axially with varices on preceding whorl. Short cord spines from IP to MP. Sinus of P1 very short. Aperture ovate. Columellar lip smooth, adherent. No parietal callus; no anal notch. Aragonitic microstructure.

Holotype. Spain• Llobregat Basin, Molins del Rei; Yellow and grey clays of Molins del Rei (quarry Anna); Pliocene (Zanclean); MNHN.F.A77751 (MNHN coll.), H: 28 mm (Figs 21A, B; 28E).

Paratypes. Spain• 2 spm; same as for the holotype; MNHN.F.A77752 (MNHN coll.) (Figs 21C, D; 28D, MNHN.F.A77753 (MNHN coll.) (Figs 21E, F; 28C).

From the Latin adjective hirtus meaning bristly, because of the cord spines of the species; gender masculine.

Yellow and grey clays of Molins del Rei (quarry Anna).

Spain, Llobregat Basin, Pliocene (Zanclean; see Salvany & Aguirre 2020), Molins del Rei (quarry Anna).

Only known from the type locality.

Protoconch not preserved. Teleoconch up to 28 mm in height (holotype), up to 7.6 mm in width, biconic in profile, composed of six whorls. Moderately low spire with carinate whorls. Last whorl up to 76% of total length. Apical angle 68°. Spiral sculpture with marked primary cords. First whorl: not preserved; from second to fourth whorl: presence of P1 and P2, P2 close to abapical suture; fifth whorl: appearance of IP; sixth whorl: appearance of abis infrasutural ramp. P1 to P5 and s1 to s3 on convex part of whorl; P6, ADP, MP and ABP on siphonal canal. Axial sculpture with high, sublamellose varices. On second whorl: 9-10 varices; on third whorl: ten varices; from fourth whorl to fifth whorl: 8-10 varices; on sixth whorl: eight varices. On spire, short cord spines on P1-P2; P1 delimiting periphery. On last whorl, short cord spines on IP, P1-P6, ADP, MP, ABP, but more developed on P1, P2 and P3; P2 delimiting periphery. Ovate aperture up to 31% of diameter and up to 74% of length of last whorl (including siphonal canal). Columellar lip, smooth, adherent. Parietal lip smooth, adherent. Outer lip lacking internal denticles. Pseudoumbilicus moderately large. Siphonal canal open, up to 41% of aperture length, slightly dorsally recurved. Aragonitic microstructure.

By its shape and its sculpture, C.(Pliocrassimurex)hirtus ‌n. subgen., n. sp. resembles members of Crassimurex ‌Merle, 1990. However, until now, Crassimurex was not known in the Pliocene and seemed extinct after the Middle Miocene. Crassimurex is currently represented by two subgenera, Crassimurex (s.s.): early Eocene to middle Eocene, France, and Crassimurex(Eopaziella): late Eocene to Middle Miocene (Europe). Therefore, the discovery of a species of Crassimurex in the Early Pliocene of Spain is surprising and extends the stratigraphic range of the genus. Crassimurex(Pliocrassimurex) differs from Crassimurex (s.s.) and C. (Eopaziella) by having a more obtuse shoulder angle. In C. (Pliocrassimurex), P2 delimits the periphery on the last whorl, whereas in Crassimurex (s.s.) and C. (Eopaziella), P1 delimits the periphery. Crassimurex (s.s.) 1990 can share cord spines with C. (Pliocrassimurex), as seen in the type species C. (s.s.)calcitrapa ‌(Lamarck, 1803) (Fig. 21G, H), but it differs by having varices which are not axially aligned. Crassimurex(Eopaziella) resembles C. (Pliocrassimurex) by having varices axially aligned but differs in lacking cord spines (see Merle et al. 2011: 178, fig. 62, pl. 144).

Shell medium sized for family, biconic. P6 not atrophied. IP, ADP, MP and ABP present. Poorly developed axial sculpture formed by slightly lamellose varices. Posterior sinus of sutural ramp shallow. Shoulder sinus of P1 reduced or missing. ID missing, D1 to D6 present. Columellar lip occasionally ornamented by 1-3 tubercles. Microstructure of three aragonite layers.

Jsowerbya ‌Merle, 2005b, Beyregrex ‌Merle ‌n. gen., and Pseudotrophonopsis ‌Merle ‌n. gen.,

This informal group includes basal taxa often erroneously attributed to the Muricopsinae (synonym of Aspellinae) because of their biconic shape and the presence of 1-3 columellar tubercles or attributed to the Trophoninae, because of the more or less lamellose varices. In the phylogenetic analysis, they represent basal taxa (Clades G, H, I, J, L of Winclada [Fig. 1; Appendix 3] and Clades 44, 43, 42, 41, 40 of PAUP [Appendix 5]). They essentially differ from members from the other subfamilies by having a combination of characters including a posterior sinus, a D6 denticle, occasional columellar tubercles, and by lacking the ID denticle. The primary cords appear late in ontogeny and IP is missing in the most basal species of this group (e.g., Beyregrex ‌sarroniensis ‌(Carez, 1879) ‌n. comb.,) from the Thanetian, whereas in the derived members (e.g., members of Jsowerbya ‌Merle, 2005b) they appear earlier, and the primary cord IP is often present. From an evolutionary point of view, members of this group are diversified from the late Paleocene to the middle Eocene with Beyregrex ‌Merle ‌n. gen., and Jsowerbya. The decline of the group begins at the Priabonian in which only Beyregrex ‌Merle ‌n. gen., and Pseudotrophonopsis ‌Merle ‌n. gen., are recorded. Beyregrex ‌Merle ‌n. gen., survives after the Eocene/Oligocene climatic crisis and is known in the Early Miocene (Burdigalian) with Beyregrex ‌cedillatus ‌(Cossmann & Peyrot, 1924) ‌n. comb.,

Murex ‌pereger ‌Beyrich, 1854, late Eocene to late Oligocene: Paris Basin, Belgium, Mainz Basin and north Germany (Fig. 22G, H).

Name based on the type species Murex ‌pereger ‌Beyrich, 1854 and using Bey (from Beyrich), the letters reg (from pereger) and rex (from Murex).

Murex ‌sarroniensis ‌Carez, 1879 [synonym: Murex ‌abbatiae ‌Wrigley, 1930], London and Paris Basins, Paleocene (Thanetian) to Early Eocene (Ypresian) (Fig. 22C, D); Murex ‌hantoniensis ‌Edwardsin Lowry et al., 1866, Hampshire Basin, England, late Eocene (Priabonian) (Fig. 22E, F); M. ‌pereger ‌Beyrich, 1854 [synonym: Murex ‌areolifer ‌Sandberger, 1861], Paris Basin, Belgium, Mainz Basin and north Germany, late Eocene (Priabonian) to late Oligocene (Chattian) (Fig. 22G, H); Hexaplex ‌brevaculeatus ‌Janssen, 1978, Paris basin, early Oligocene (Rupelian), Mainz basins, early Oliogocene and North Germany, late Oligocene; Murex(Poirieria)cedillatus ‌Cossmann & Peyrot, 1924, Aquitaine Basin, France, Early Miocene (Burdigalian).

Biconic teleoconch with subangular whorls. D1 well developed. P6 not atrophied. IP weak, ADP, MP present. Varices formed by 1-3 lamellae abaxially turned on the early whorls, occasionally reinforced by several lamellae on the succeeding whorls. P1 spine short. Shoulder sinus widely open, short, moderately deep. Posterior sinus on sutural ramp shallow. Columellar lip usually smooth.

Members of Beyregrex ‌Merle ‌n. gen., differ from those of Jsowerbya in having a deeper shoulder sinus, more lamellose varices, and a smooth columellar lip. Pseudotrophonopsis ‌Merle ‌n. gen., is also distinguishing from Beyregrex ‌Merle ‌n. gen., in having rounded whorls, strong secondary cords of equal strength to the primary cords, and numerous split internal denticles within the outer lip (D1 to D5). Represented by five species, Beyregrex ‌Merle ‌n. gen., is the most diverse genus of the group. It extends from the Anglo-Parisian late Paleocene/early Eocene with B. sarroniensis ‌(Carez, 1879) ‌n. comb., to the Early Miocene (Burdigalian) from the Aquitaine Basin with B. cedillatus ‌(Cossmann & Peyrot, 1924). Beyregrex ‌sarroniensis ‌n. comb. displays several primitive characters, such as missing IP, MP and ABP. During the middle Eocene, no Beyregrex ‌Merle ‌n. gen., species are known, but the genus re-emerges in the late Eocene of Hampshire with B. hantoniensis (Edwards in Lowry et al., 1866), n. comb.. That species is closely related to B. pereger ‌(Beyrich, 1854) ‌n. comb., occurring in the Paris Basin and in Germany (Mainz Basin and north Germany) during the Oligocene. According to Janssen (1978), B. brevaculeatus ‌(Janssen, 1978) is another species. It was originally described from the Chattian of Germany (at Glimmerode) but is also recorded in the Rupelian of the Paris Basin (Merle 1999; Lozouet 2012). For the Neogene, B. cedillatus ‌(Cossmann & Peyrot, 1924) from the Burdigalian of the Aquitaine Basin demonstrates that the genus was not restricted to northern Europe.

Buccinum ‌defossum ‌Pilkington, 1804, middle Eocene (Bartonian) of England (Hampshire) (Fig. 22I-K), and late Eocene (Priabonian) of France (Paris Basin) (Fig. 22L).

Name based on the Greek prefix pseudo (false), because the superficial resemblance of the type species with members of the genus Trophonopsis ‌Bucquoy & Dautzenberg, 1882.

Buccinum ‌defossum ‌Pilkington, 1804 [synomym: Fusus ‌sublamellosus ‌Deshayes, 1835], England, Hampshire Basin, middle Eocene (Bartonian) and France, Paris Basin, late Eocene (Priabonian).

Smooth multispiral protoconch (Fig. 22K). Biconic teleoconch with rounded whorls. P6 not atrophied. IP, ADP, MP, ADP present. Primary P1 to P6 present on first whorl (Fig. 22K). Coarse secondary cords equal in strength to primary cords. Varices low, formed by one short abaxial lamella. No P1 spine. No shoulder sinus. Posterior sinus shallow. Columellar lip smooth. Internal denticles D1 to D6 split. ID missing.

This strange species differs from members of Jsowerbya ‌Merle, 2005b and Beyregrex ‌Merle ‌n. gen., by having rounded whorls, no shoulder sinus and by having strongly developed secondary cords. Because of its rounded whorls, it resembles members of Nucellopsis, but they are distinguishing by their buccinoid shape and by a later appearance of the primary cords. The phylogenetic analysis (Winclada tree, Fig. 1) suggests that it could represent a stem group of Jsowerbya. We therefore propose the new genus Pseudotrophonopsis ‌Merle ‌n. gen., as P. ‌defossa ‌(Pilkington, 1804) ‌n. comb., cannot be attributed to Jsowerbya or Beyregrex ‌Merle ‌n. gen., Pseudotrophonopsis ‌defossa ‌n. comb. occurs in the Bartonian from Hampshire (England) and is known in the Priabonian from the Paris Basin, where it was identified as Fusus ‌sublamellosus ‌Deshayes, 1835. The protoconch of the English and French populations (Fig. 22K) is similar and the teleoconch of the French population is only less thick (personal observation DM). Therefore, we believe that they belong to the same species. Glibert (1963) recorded this species in the Bartonian from the Paris Basin, but we cannot confirm this record based on material in the examined collections (MNHN and private collections). It is possible that this species was mistaken for Jsowerbya ‌depauperata ‌(Deshayes, 1865).

Nucellopsis ‌Merle, 2005 by present designation.

Nucellopsis ‌Merle, 2005 and Pronucellopsis ‌n. gen.

Shell medium sized for family. Bucciniform shape with rounded whorls. P6 not atrophied. ADP, MP present; ABP and IP present in some species of Nucellopsis. Poorly developed axial sculpture formed by weakly lamellose varices. Posterior sinus on sutural ramp deep. D1 to D6 present. ID missing. Columellar lip usually smooth. Microstructure of three aragonite layers.

The Nucellopsinae ‌Merle ‌n. subfam., includes basal taxa often erroneously attributed to the Muricopsinae (synonym of Aspellinae) because of their biconic shape or attributed to the Trophoninae because of their weakly lamellose varices. The Nucellopsinae ‌Merle ‌n. subfam., (Clade D of Winclada or Clade 47 of PAUP) includes Paleocene and Eocene species that usually share a late appearance of the primary cords (P1 to MP), rounded whorls and buccinoid shape in Nucellopsis.

Murex ‌plicatilis ‌Deshayes, 1835 by original designation. Early Eocene: Paris Basin (France) and London Basin (England).

Holotype. France• Paris Basin, Val d’Oise, Saint-Witz (Le Guépelle); Sable du Guépelle (biozone NP16); middle Eocene (early Bartonian, Auversian); MNHN.F.R64955 (Merle coll.), figured specimen in Merle (2005c: fig. 2H), H: 21.5 mm (Figs 23E, F; 28F).

Paratypes. France• 1 spm; same as for the holotype; MNHN.F.A71797 (Pons coll.), figured specimen in Merle (2005c: fig. 1G) • 1 spm; idem; MNHN.F.A71798 (Pons coll.), figured specimen in Merle (2005c: fig. 1H, I)•1 spm; idem; MNHN.F.J02585 (Cossmann coll.); figured in Cossmann & Pissarro (1911: pl. 36 fig. 169bis-2)•1 spm; idem; MNHN.F.A90963 (Faullummel coll.) (Figs 23C, D; 28G)•1 spm; idem; MNHN.F.A90964 (Faullummel coll.) (Fig. 23A, B)•6 spm; idem; MNHN.F.A90962 (Faullummel coll.).

France • 1 spm; Paris Basin, Val d’Oise, Saint-Witz (Le Guépelle); Sable du Guépelle (biozone NP16); early Bartonian; MNHN.F.A27771 (Pacaud coll.)•1 spm; Paris Basin, Val d’Oise, Auvers-sur-Oise (Bois-le-Roi); idem; MNHN.F.A27770 (Pacaud coll.)•1 spm; Paris Basin, Aisne, Nogent-l’Artaud; idem; MNHN.F.A90565 (Margerie coll.)•1 spm; Paris Basin, Aisne, Bézu-le-Guéry; idem; MNHN.F.A90825 (Faullummel coll.)•1 spm; Paris Basin, Aisne, Mont-Saint-Martin; idem; MNHN.F.J07787 (UMPC coll.)•1 spm; Paris Basin, Seine-et-Marne, Sainte-Aulde (Caumont); idem; MNHN.F.A90826 – (Faullummel coll.)•1 spm; Paris Basin, Seine-et-Marne, Luzancy; idem; MNHN.F.A90827 (Faullummel coll.)•Paris Basin, Oise, Loisy; idem; MNHN.F.A90828 (Faullummel coll.).

From the Latin adjective parisiensis, the geographic origin of this species being the Paris Basin.

Sable du Guépelle (biozone NP16, see Gély & Lorenz (1991: pl. 1), middle Eocene (early Bartonian, Auversian).

France, Paris Basin, Val d’Oise, Saint-Witz (Le Guépelle), INPN (2024).

Paris Basin (France), middle Eocene (early Bartonian, Auversian).

Smooth, multispiral protoconch of up to 2.5 whorls (see Merle 1999: pl. 23, fig. 5, holotype). Biconic teleoconch, up to 21.5 mm in height, up to 14 mm in width, biconic in profile and composed of six whorls. Moderately high spire with rounded or subcarinate whorls. Last whorl up to 73% of total length. Apical angle 64°. Spiral sculpture with strongly marked primary cords. First whorl: appearance of P1, s1, P2; P1 less marked than s1-P2; from second to third whorl, no addition of cord, P1 of equal strength to s1-P2; on fourth whorl, appearance of IP poorly marked. Fifth whorl: no addition of cord. Last whorl: IP (weak), P1 to P6, P6 not atrophied (on convex part of whorl), ADP, MP and ABP; s1 to s3 dominant, equal in strength to primary cords. First to second whorl: twelve varices. Third to fourth whorl: eleven varices. Fifth whorl: ten varices. Sixth whorl: nine varices. Varices more or less sublamellose. No true spine except angular point on P1. Suboval aperture up to 31% of diameter and up to 66% of length of last whorl (including siphonal canal). Columella slightly angular. Columellar lip narrow, adherent, smooth or with one tubercle. Outer lip with five or six denticles including D1 to D6. D1 appearing late in ontogeny. D6 occasional. ID missing. Siphonal canal, short, open, up to 31% of aperture length, slightly dorsally recurved. Pseudo-umbilicus deep, rather wide.

Nucellopsis ‌ponsi ‌Merle, 2005c differs from N. ‌parisiensis ‌n. sp., N. plicatilis ‌(Deshayes, 1835) andN. ‌dudariensis ‌(Strauzs, 1966) by having a deeper posterior sinus and weaker spiral cords. Its columellar lip is occasionally ornamented by a weak denticle mid-columella, whereas it is usually smooth in the other three species. Nucellopsis ‌parisiensis ‌n. sp., N. plicatilis and N. ‌dudariensis bear more numerous cords on the siphonal canal (MP and ABP) and more secondary cords than N. ‌ponsi. Nucellopsis ‌parisiensis ‌n. sp. differs from N. plicatilis by the appearance of P1 and s1 on the first teleoconch whorls, by having more angular teleoconch whorls, and by usually lacking D6. Both species share a multispiral protoconch. Nucellopsis ‌parisiensis ‌n. sp. was erroneously identified as Jsowerbya ‌depauperata ‌(Deshayes, 1865) by Cossmann & Pissarro (1911) in spite of numerous differences, such as a narrower shell shape, IP on the sutural ramp and a deeper anal sulcus (Merle 2005b). Merle (1999, 2005c) identified N. ‌parisiensis ‌n. sp. as N. ‌dudariensis due to several similarities with the holotype of Cantharus(Pollia)dudariensis(Fig. 23G, H) from the upper Lutetian of Hungary [apical angle near 64°, subangular whorls, thick varices, poorly developed shoulder sinus, five internal denticles (D1 to D5, D6 missing), angular columella with one small tubercle]. These similarities suggest that N. ‌parisiensis ‌n. sp. and N. dudariensis are closely related. However, close examinaton of the holotype of N. dudariensis (Fig. 23G, H) reveals several differences with the Parisian species, such as the late appearance of s1 on the last whorl, s1 less coarse than the primary cords P1 and P2, fewer secondary cords (only s1 instead s1 to s3), and the stronger development of IP.

Pronucellopsis ‌pacaudi ‌Merle, Ledon & Goret ‌n. sp., by present designation.

Pronucellopsis ‌pacaudi ‌Merle, Ledon & Goret ‌n. gen., n. sp.,, Paris Basin, France; Paleocene (Thanetian).

Subfusiform teleoconch with rounded whorls. Poorly marked cords. No IP. P6 weak, not atrophied. ADP, MP weak. Secondary cords (s1, s2, s3, s4) appearing late in ontogeny. Primary cords P3 to P6 appearing after third whorl. Varices formed by one or two abaxial lamellae on early whorls, occasionally reinforced by several lamellae on succeeding whorls. Number of varices: between 13 and 15. Weak, rounded shoulder sinus. Posterior sinus on sutural ramp deep. Internal denticles usually weak: ID missing. D1 stronger than D2. D6 present. Columellar lip smooth. Siphonal canal narrow. Microstructure of three aragonite layers.

Pronucellopsis ‌Merle ‌n. gen., is the sister taxon of Nucellopsis (clade E, Fig. 1). Both taxa share a weak or missing shoulder sinus. Pronucellopsis ‌Merle ‌n. gen., is easily differentiated in having a more elongated shape and poorly developed internal denticles. It also displays poor development of the spiral cords which always remain weak and appear late in ontogeny. This feature is present in early muricids from the Paleocene (e.g., Poirieria or Flexopteron) and seems a plesiomorphic character.

Holotype. France • Paris Basin, Marne, Châlons-sur-Vesle; Sables de Châlons-sur-Vesle, biozone NP9; Paleocene (Thanetian); MNHN.F.J02587 (Cossmann coll.), figured specimen in Cossmann & Pissarro (1911: pl. 36, fig. 169bis-5), H: 22 mm (Figs 24A, B; 28H).

Paratypes. France • 3 spm; same as for the holotype; MNHN.F.A90831 (Pacaud coll.)•2 spm; idem; MNHN.F.A90832 (MNHN coll.)•7 spm; Paris Basin, Marne, Jonchery-sur-Vesle; Paleocene (Thanetian); MNHN.F.A90833 (Staadt coll.)•1 spm; idem; MNHN.F.A90829 (Pacaud coll.) (Figs 24C, D; 28I)•8 spm; idem; MNHN.F.A90830 (Pacaud coll.)•6 spm; Paris Basin, Oise, Abbecourt (Bois-des-Godins), Paleocene (Thanetian); MNHN.F.A90834 (Pacaud coll.).

1 spm; Paris Basin, Marne, Jonchery-sur-Vesle; Paleocene (Thanetian); UPMC (Munier-Chlamas coll.), figured specimen of Merle (1999: pl. 66, b-d) (Fig. 24E, F)•1 spm; idem; MNHN-IM (Staadt coll.), figured specimen of Merle (1999: pl. 66, c-e) (Fig. 24G, H)•5 spm; idem; MNHN.F.A91007 (Staadt coll.).

Dedicated to Jean-Michel Pacaud (MNHN) who has for a long time observed that Murex ‌multistriatus ‌Deshayes, 1865 is a junior synonym of Fusus ‌angusticostatus ‌Melleville, 1843 (now Lyrofusus ‌angusticostatus).

Sables de Châlons-sur-Vesle, biozone NP9, see Huyghe et al. (2015: fig. 2), Paleocene (Thanetian).

France, Paris Basin, Marne, Châlons-sur-Vesles, see Fritel (1910: 77).

Paris Basin (France), Paleocene (Thanetian), Sables de Bracheux and Sables de Châlons-sur-Vesle.

Unknown protoconch. Teleoconch, up to 22 mm in height, up to 12.5 mm in width, subfusiform in profile, composed of seven whorls. High spire with rounded whorls. Last whorl up to 67% of total length. Apical angle 45°. Spiral sculpture with weak primary cords. First and second whorls: no spiral scultpure; third and fourth whorls, P1 weak. Fifth whorl: appearance of P2. Sixth whorl: appearance of s1. Last whorl and seventh whorl: P1 to P6, s1, s2, s3 and s4 (convex part of whorls), ADP, MP (siphonal canal). Axial sculpture on first to second whorl: not observed; third whorl: nine varices; fourth whorl: 14 varices; fifth whorl: 14-16 varices; sixth whorl: 11-19 varices; seventh whorl: 12-23. Varices finely lamellose. No spine on P1. Wide, ovate aperture up to 45% of diameter and up to 84% of length of last whorl (including siphonal canal). Posterior sinus deep, no shoulder sinus. Columella slightly angular. Columellar lip smooth, narrow and adherent. Outer lip with occasional denticles. D1 to D6 small, poorly developed, ID missing. Siphonal canal, short, open, up to 23% of aperture length, slightly dorsally recurved. Pseudoumbilicus narrow.

Deshayes (1865) described Murex ‌multistriatus and illustrated a specimen bearing a buccinid shape (pl. 88, fig. 810). In fact, this species is a junior synonym of Fusus ‌angusticostatus ‌Melleville, 1843 (now Lyrofusus ‌angusticostatus). However, Cossmann (1889) and Cossmann & Pissarro (1911) confused this buccinoid species with a Thanetian Muricidae, which they mistakenly called Murex ‌multistriatus. We described this species as Pronucellopsis ‌pacaudi ‌Merle, Ledon & Goret ‌n. gen., n. sp., It can be compared to Nucellopsis ‌ponsi ‌Merle, 2005c or N. plicatilis ‌(Deshayes, 1835) from the late Paleocene-early Eocene of the Paris Basin. Juveniles of P. ‌pacaudi ‌Merle, Ledon & Goret ‌n. gen., n. sp., share with these Nucellopsis a buccinid shape, but at the end of its growth P. ‌pacaudi ‌Merle, Ledon & Goret ‌n. gen., n. sp., displays a subfusiform shape with a high spire. The sculptures (both axial and spiral) of P. ‌pacaudi ‌Merle, Ledon & Goret ‌n. gen., n. sp., and the internal denticles of outer lip are also less developed. Because of its high spire, P. ‌pacaudi ‌Merle, Ledon & Goret ‌n. gen., n. sp., resembles some elongated Beyregrex ‌sarroniensis ‌(Carez, 1879) ‌n. comb., but in B. ‌sarroniensis ‌n. comb. the spiral sculpture and the internal denticles are always more strongly developed. In addition, P1 spine is present in B. ‌sarroniensis ‌n. comb., whereas it is missing in P. ‌pacaudi ‌Merle, Ledon & Goret ‌n. gen., n. sp.,