The Lower Pliocene marine gastropods of Santa Maria Island, Azores: Taxonomy and palaeobiogeographic implications Author Sacchetti, Claudia 0000-0002-3225-3139 claudiasacc@icloud.com Author Landau, Bernard 0000-0002-7768-8494 bernardmlandau@gmail.com Author Ávila, Sérgio P. 0000-0002-3225-3139 claudiasacc@icloud.com text Zootaxa 2023 2023-05-24 5295 1 1 150 http://dx.doi.org/10.11646/zootaxa.5295.1.1 journal article 53396 10.11646/zootaxa.5295.1.1 82286fdc-a858-447c-9980-da2e8985d19c 1175-5326 7965273 F3A52660-70B8-439F-A7A0-F45ADC975EA5 Crassopleura maravignae (Bivona, 1838) Plate 6 C, D Pleurotoma elegans Scacchi 1836: 43 , pl. 1, fig. 8 ( non Defrance, 1826). Pleurotoma incrassata Dujardin 1837: 292 , pl. 20, fig. 28 ( non G.B. Sowerby, I 1834 ). * Pleurotoma Maravignae Bivona, Ant. In Bivona And. 1838: 13, pl. 1, fig. 12. Pleurotoma Maravignae Biv. — Calcara 1839: 8 , pl. 1, fig. 3. Pleurotoma elegans Scacchi — Calcara 1841: 54 . Pleurotoma elegans Scacchi — Philippi 1844: 168 , pl. 26, fig. 5. Raphitoma incrassata Duj. — Bellardi 1847: 108 , pl. 4, fig. 27. Pleurotoma subincrassata d’Orbigny 1852: 62 ( nom. nov. pro. P. incrassata Dujardin, 1837 , non G.B. Sowerby, I 1834 ). Pleurotoma incrassata Duj. —Ĥrnes 1853: 383, pl. 40, fig. 14. Pleurotoma Hybrida Millet 1854: 161 ( nomen nudum ). Pleurotoma incrassatum Duj. — Brugnone 1862: 20 , pl. 1, fig. 12. Pleurotoma perturrita Bronn in Reiss 1862: 29 , pl. 1, fig. 9. Pleurotoma perturrita Bronn — Mayer 1864: 68 , pl. 6, fig. 50. Pleurotoma hybrid Millet 1865: 161 ( non Grateloup, 1845 ). Drillia incrassate (Duj.) — Bellardi 1877: 140 , pl. 5, fig. 1. Drillia incrassate var. Rhodanica Fontannes 1879: 46 , pl. 4, fig. 10. Drillia ( Cymatosyrinx ) incrassata var. acutespirata Sacco 1904: 46 . Drillia ( Cymatosyrinx ) incrassata var. magnocostulata Sacco 1904: 47 . Drillia ( Cymatosyrinx ) incrassata var. miominor Sacco 1904: 47 , pl. 12, figs. 47, 48. Drillia ( Cymatosyrinx ) incrassata var. dertomagna Sacco 1904: 47 . Drillia ( Cymatosyrinx ) incrassata Dujardin — Cipolla 1914: 122 , pl. 12, fig. 10. Drillia ( Cymatosyrinx ) incrassata mut. Maravignae Biv. – Cipolla 1914: 123 , pl. 12, fig. 11. Drillia ( Cymatosyrinx ) incrassata var. miominor Sacco – Cipolla 1914: 123 , pl. 12, fig. 12. Drillia incrassate (Dujardin) —Harmer ( partim ) 1915: 222, pl. 27, fig. 28. Drillia ( Cymatosyrinx ) incrassata var. minor Montanaro 1937: 159 , pl. 7, figs. 41-43. Drillia ( Cymatosyrinx ) subincrassata Peyrot 1938: 275 . Clavus ( Crassopleura ) maravignae (Bivona) —Wenz 1944: 1397, fig. 3948. Clavus ( Crassopleura ) maravignae Bivona, 1838 — Glibert 1954: 33 , pl. 1, fig. 12, pl. 5, fig. 4. Clavus ( Crassopleura ) maravignae Bivona, 1838 — Brébion 1964: 553 . Crassopleura maravignae (Bivona, 1838) — Powell 1966: 93 , pl. 14, fig. 12. Crassopleura maravignae (Bivona, 1938 [ sic ])— Nordsieck 1977: 11 , pl. 1, fig. 3. Crassopleura incrassata ( Dujardin, 1837 ) —Bernasconi & Robba 1984: 270, pl. 2, fig. 1. Crassopleura maravignae (Bivona, 1838) — Van Aartsen et al . 1984: 43 , fig. 207. Crassopleura incrassata ( Dujardin, 1837 ) — Vera-Pelàez 1996: 371 , text-figs. 16a-d, 19a-d, 21a, b, 26, pl. 25, figs. 1-12. Crassopleura incrassata ( Dujardin, 1837 ) — Chirli 1997: 43 , pl. 12, figs. 2-5. Crassopleura maravignae (Bivona, 1838) — Ardovini & Cossignani 1999: 67 , 68, unnumbered fig top row right. Crassopleura maravignae (Bivona Ant. in Bivona And., 1838)— Ávila et al . 2000: 153 . Crassopleura incrassata ( Dujardin, 1837 ) —Vera-Pelàez 2002: 197, pl. 3, figs. D, E. F, pl. 11, figs. I, J. Crassopleura maravignae (Bivona Ant. In Bivona And., 1838)— Scarponi & Della Bella 2003: 21 , figs. 2, 3, 14. Crassopleura maravignae Bivona Ant. in Bivona And., 1838—Ávila 2005: 118, 256. Crassopleura maravignae (Bivona, 1838) — Martins et al . 2009: 65 : 98 , pl. xv, figs. 248-250. Crassopleura maravignae (Bivona Ant. In Bivona And., 1838)— Sosso & Dell’Angelo 2010: 44 : 60 unnumbered fig. bottom left. Crassopleura maravignae (Bivona Ant. In Bivona And., 1838)— Landau et al . 2011: 36 , pl. 18, fig. 12. Crassopleura maravignae (Bivona Ant. in Bivona And., 1838)—Cordeiro et al . 2015: 870. Crassopleura maravignae (Bivona Ant. In Bivona And., 1838)—Landau & Harzhauser 2023: xx, pl. xx, fig. xx. non Pleurotoma incrassata Duj. — Nyst 1878 , pl. 3, fig. 10 [= Elaeocyma nysti (Harmer, 1915) ]. non Pleurotoma incrassata Duj. — Nyst 1882: 45 [= Elaeocyma nysti (Harmer, 1915) ]. non Drillia incrassata var. dertomagna Sacco —Harmer 1915: 223, pl. 27, fig. 29 [= Elaeocyma nysti (Harmer, 1915) ]. non Drillia incrassata var. miominor Sacco —Harmer 1915: 223, pl. 27, fig. 30 [= Elaeocyma nysti (Harmer, 1915) ]. non Drillia incrassata var. nysti Harmer 1915: 224 , pl. 27, fig. 31 [= Elaeocyma nysti (Harmer, 1915) ]. non Drillia incrassata var. crassa (A. Bell) —Harmer 1915: 224, pl. 27, fig. 32 [= Elaeocyma nysti (Harmer, 1915) ]. non Drillia incrassata Duj. — Csepreghy-Meznerics 1953: 8 , pl. 1, figs. 5, 6 [ = ‘Crassopleura’ sigmoidea ( Bronn, 1831 ) ]. Santa Maria material. Maximum height 12.0 mm, width 4.5 mm. Single specimen from Pinheiros, Santa Maria Island, Touril Complex, Lower Pliocene. Description. (Based on Estepona Basin specimens, as no specimens from Santa Maria are available). Shell small to medium-sized, solid, fusiform, with tall, conical spire. Protoconch multispiral, conical, of 3.5 smooth, convex whorls, with small nucleus. Junction with teleoconch sharply delimited by sinusigera. Teleoconch of up to seven whorls with almost vertical, weakly concave subsutural ramp, poorly delimited by weak shoulder, convex below, separated by superficial, undulating suture. Axial sculpture of narrow, opisthocline sinuous ribs, 14-19 on last whorl, narrow over subsutural ramp, slightly swollen towards lower suture. Spiral sculpture restricted to 3 or 4 flattened cords over siphonal fasciole. Last whorl subsutural ramp slightly concave, broadly and weakly rounded below, not constricted at base, ribs weakening at periphery, extending over base in most specimens; base not delimited; siphonal fasciole weakly delimited, flattened, bearing broad spiral cords. Aperture ovate; outer lip strongly thickened by broad labial varix; anal sinus shallow, symmetrically U-shaped, occupying entire subsutural ramp, with apex mid-ramp; siphonal canal very short and broad, straight, unnotched. Columella straight, smooth. Columellar and parietal callus thickened, sharply delimited, forming moderately broad callus rim; well-developed tuberculose parietal pad developed adapically delimiting medial border of anal canal. Colour patten is preserved consisting of a single narrow orange horizontal stripe running at about two-thirds whorl height. Discussion. As discussed by Scarponi & Della Bella (2003: 22) , the correct name for this species is Crassopleura maravignae (Bivona 1838) , Dujardin’s (1837) name Pleurotoma incrassata being a primary homonym of P. incrassata G. B. Sowerby I, 1834. Bronn in Reiss (1862: 29) described Pleurotoma perturrita Bronn in Reiss, 1862 based on a single specimen from Pinheiros. Judging from the original illustration, this species is almost certainly synonymous with Crassopleura maravignae (Bivona 1838) , and Bronn’s taxon is considered a junior subjective synonym herein. Some specimens with fewer ribs resemble specimens of Crassopleura sigmoidea ( Bronn, 1831 ) , but can be differentiated in that the anal sinus is evenly U-shaped and does not produce a notch on the parietal wall. Scarponi & Della Bella (2003) considered this difference to be of generic significance and placed ‘ C. ’ sigmoidea species in the genus with exclamation marks. Crassopleura maravignae (Bivona 1838) is widespread from the Middle Miocene to present-day European assemblages, although there are small differences between populations. Glibert (1954: 33) commented that the middle Miocene specimens from the Loire Basin were slightly slenderer than those from the Holocene Mediterranean and that the ribs were less serrated and more oblique. The Estepona specimens have 14-19 ribs on the last whorl. Scarponi & Della Bella (2003: 23) noted that the most common Mediterranean Pliocene form had a similar number of ribs (14-18), and Landau et al . (2020c) counted 15-20 ribs for the upper Miocene specimens from northwestern France , all slightly fewer than present-day specimens (21-25). However, the older Atlantic Middle Miocene population from the Loire Basin had anything between 15 and 30 axial ribs per whorl ( Glibert 1954: 33 ). The colour pattern preserved in the Estepona specimens is also seen in some, but not all, extant shells. Some references to Crassopleura maravignae (as Pleurotoma or Drillia incrassata ) ( Nyst 1878 , 1882 ; Harmer 1915) do not correspond to this species, but to Elaeocyma nysti (Harmer, 1915) ( Marquet 1998a , b). Distribution. Middle Miocene: Atlantic (Langhian), Loire Basin, France ( Dujardin 1837 ; Glibert 1954 ); Paratethys, Austria (Ĥrnes 1853); Proto-Mediterranean, Italy ( Bellardi 1877 ). Upper Miocene (Tortonian and Messinian): Atlantic, NW France ( Millet 1854 , 1865 ; Glibert 1954 ; Brébion 1964 ; Landau et al . 2020c ); Proto-Mediterranean, Italy ( Bellardi 1877 ; Montanaro 1937 ). Lower Pliocene:?NSB, Coralline Crag, England (Harmer 1915); Atlantic, Santa Maria Island (Bronn in Reiss 1862 ; Mayer 1864 ), Guadalquivir Basin, S. Spain ( Landau et al . 2011 ); western Mediterranean, NE Spain ( Gili & Martinell 1993 ), S. France ( Fontannes 1879 ); central Mediterranean, Italy ( Chirli 1997 ; Scarponi & Della Bella 2003 ; Sosso & Dell’Angelo 2010 ). Upper Pliocene: western Mediterranean, Estepona Basin, S. Spain ( Vera-Pelàez 1996 , 2002; Landau & Harzhauser 2023), central Mediterranean, Italy ( Calcara 1839 , 1841 ; Bellardi 1877 ; Cipolla 1914 ; Bernasconi & Robba 1984). Present-day: Eastern Atlantic frontage from Portugal south to NW Africa, into the Mediterranean ( Nordsieck 1977 ), central Mediterranean ( Ardovini & Cossignani 1999 ) and the Azores , Madeira , and Canary Islands.