Genus Clavatula Lamarck, 1801

Type species. Clavatula coronata Lamarck, 1801 [= Turris regia Röding, 1798], by monotypy (Lamarck 1801: 84). Present-day, tropical Eastern Atlantic.

Diagnosis. “ Shell moderately large to large sized 15–55 mm, very solid, buccinoid, coarsely axially and spirally sculptured; with a tall, often coronated spire, and a truncated body-whorl, terminated in a relatively short reflected and deeply notched anterior canal, which is backed by a prominent ridge-margined fasciole, encircling a false umbilical cavity that can be quite wide. Protoconch relatively small 2–3 smooth whorls. Adult sculpture of strong but irregular spiral cords which may be smooth or granose, often produced into nodules or pointed up-curved spines… The suture, typically, is at the top of a wide broadly rounded fold which clasps high on the preceding whorl and half envelopes the peripheral carina ” (Powell 1966: 57). In addition, we consider a bipartite sculpture on early teleoconch whorls with a weak, smooth adapical subsutural cord and a prominent suprasutural cord bearing large beads as diagnostic.

Discussion. When establishing Clavatula as a new genus, Lamarck (1801: 84) referred to Murex Turris coronata described by Chemnitz in Martini & Chemnitz (1795: 114, pl. 190, figs 1831, 1832) from Guinea. This work, however, is placed by the ICZN on the Official Index of Rejected Works. By referring to the illustration in Chemnitz in Martini & Chemnitz (1795), Lamarck (1801) made the name Clavatula coronata available. However, Chemnitz in Martini & Chemnitz (1795: 114) stated that he had already described and illustrated the same species as Turris babylonica coronata in Chemnitz in Martini & Chemnitz (1780: 176, vignette 39, fig C). This name is not available either. However, Röding (1798: 124) explicitly referred to “Martini 4. p. I43. ving. 38 Lt. C” in Chemnitz in Martini & Chemnitz (1780) and provided the name T [urris]. Regia for that species, and listed three specimens (Cernohorsky 1974: 185). Therefore, Turris regia Röding, 1798 has priority over Clavatula coronata Lamarck, 1801. This has been overlooked by subsequent authors such as Mörch (1852: 72) and Tryon (1884: 229), who discussed Clavatula muricata (Lamarck, 1822) as junior synonym of Clavatula coronata Lamarck, 1801 (see also Tucker 2004: 244). The first to clarify this confusion was Juan Horro (Vigo, Spain, pers. comm, March 2021), who kindly shared this information with us.

The specimens illustrated by Chemnitz in Martini & Chemnitz (1780) were part of the kongelige danske Kunstkammer (the Royal Danish Kunstkammer) and are stored in the Natural History Museum of Denmark in Copenhagen. As Chemnitz in Martini & Chemnitz (1795) himself noted, the illustrations in his book from 1780 (vignette 39, fig C) are poor, making it impossible to identify the illustrated specimen. Cernohorsky (1974) mentioned three specimens as syntypes of Clavatula regia. Originally, the lot described by Chemnitz in Martini & Chemnitz (1780) consisted of four specimens, which were illustrated by Gundestrup (1991) and are still stored in the collection (pers. comm. Tom Schiøtte, Natural History Museum of Denmark). To stabilize the complex taxonomic history of this species we select specimen Fig. 8F as lectotype of Clavatula regia (Röding, 1798) (NHMD-916083). The specimens illustrated as Fig. 8E (NHMD-230855), Fig. 8G (NHMD-230856) and Fig. 8H (NHMD-90802) are paralectotypes.

Cernohorsky (1974) treated Clavatula muricata (Lamarck, 1822) as subjective junior synonym of Clavatula regia. Molecular data by Nicolas Puillandre (pers. comm., October 2021) do not contradict this view, but more samples would be needed to evaluate the status of both species and we have not studied the type specimens of Clavatula muricata.

Distribution and stratigraphy. Clavatula sensu stricto occurs along the coast of West Africa (Strebel 1912; Knudsen 1952; Bernard 1984). The fossil record is difficult to evaluate due to the poor information on early teleoconch sculptures. The oldest record of Clavatula s.s. is Clavatula sorini nov. sp. from the Langhian of the Central Paratethys. All other records are treated in the following as ‘ Clavatula ’ sensu latu and most probably should be excluded from Clavatula.

Paleoenvironment. Clavatula sensu stricto is documented from inner neritc environments down to few tens of meters of waterdepth (Bernard 1984).