Late Miocene Conidae (Mollusca: Gastropoda) of Crete (Greece). Part 1: genera Conilithes Swainson, 1840 and Conus (Kalloconus) da Motta, 1991 Author Psarras, Christos National and Kapodistrian University of Athens, Faculty of Geology & Geoenvironment, Department of Hist. Geology-Paleontology, Athens (Greece) cpsarras@geol.uoa.gr Author Koskeridou, Efterpi National and Kapodistrian University of Athens, Faculty of Geology & Geoenvironment, Department of Hist. Geology-Paleontology, Athens (Greece) ekosker@geol.uoa.gr Author Merle, Didier CR 2 P (CNRS, MNHN, UPMC, Sorbonne Université), Département Origines et Évolution, Muséum national d’Histoire naturelle, case postale 38, 57 rue Cuvier, F- 75231 Paris cedex 05 (France) didier.merle@mnhn.fr text Geodiversitas 2021 2021-12-02 43 24 1309 1339 journal article 3058 10.5252/geodiversitas2021v43a24 d9b664cc-3197-44d5-92ca-08f8ce0fcc4f 1638-9395 5764710 urn:lsid:zoobank.org:pub:D36D1E14-73BE-4176-8024-F3673A65B8C1 Conus ( Kalloconus ) asterousiaensis n. sp ( Figs 24 , 25 , 26 , 27 ; Table 11 ) urn:lsid:zoobank.org:act: CD6E9758-6E97-45B5-A8F0-1C900A4417A7 FIG. 22. — Conus ( Kalloconus ) letkesensis ( Harzhauser & Landau, 2016 ) from Filippi, Crete (Greece). Specimen AMPG(IV) 2682 with small dense dashes and irregular flammulae on the spire. Specimen shown under natural light ( A1-A2 , A4 ) and UV light. Scale bar: 1 cm. Conus raristriatus – Davoli 1972: 74 , pl. 8, fig. 1a, b. DIAGNOSIS. — Conus ( Kalloconus ) of medium-large size, robust shell with conical spire, with colour pattern of spiral rows of quadrangular dots and fluorescent bands on spire whorls. TYPE MATERIAL. — Holotype : AMPG(IV) 2683 , Filippi ; 1 paratype AMPG(IV) 2688 , Filippi ; 1 paratype AMPG(IV) 2689 , Tefeli ; 1 paratype MNHN.F. A72644 , Crete . TYPE LOCALITY. — Filippi, 35°02’07.2”N , 25°15’00.5”E , Messara Basin, Tortonian , Crete , Greece . STRATIGRAPHIC RANGE. — Tortonian of Greece (Messara Basin, Crete), and Italy. ETYMOLOGY. — Name taken from the Asterousia Mountain range south of the locality, Crete , Greece . OTHER MATERIAL EXAMINED. — Greece . Filippi : four specimens AMPG(IV) 2684-2687 ; Partira : one specimen MNHN.F. A72650 ; Tefeli : one specimen AMPG(IV) 2690 ; Crete : seven specimens MNHN.F. A72642 to MNHN.F. A72643 and MNHN.F. A72645 to MNHN.F. A72649 . All specimens display a colour pattern under UV light . FIG. 23. — Pattern of the dashes on Conus ( Kalloconus ) letkesensis ( Harzhauser & Landau, 2016 ) . Dashed colour pattern is from AMPG(IV) 2682 (Fig. 22) from Filippi, Crete (Greece). Scale bar: 1 cm. SHELL DESCRIPTION Medium-sized, stout shell. Spire straight to coeloconoid, highly conical to flat. Protoconch multispiral ( Fig. 25B ). Early spire whorls elevated, coeloconoid, smooth, with straight to convex outline. Later spire whorls straight to concave, with smooth, conical outline. Last spire whorl convex, slightly striate. Shoulder rounded to angulated. Maximum diameter just below shoulder. Subsutural flexure shallow, moderately to weakly curved, strongly asymmetrical. Last whorl conical, straight. Aperture narrow, straight, widening near siphonal canal.Siphonal canal wide, short. Fasciole short, twisted. There are two extremes of a form. The first extreme form (form 1, Fig. 27A, G ) consists of conical spire whorls of medium height and rounded shoulder. The second extreme form (form 2, Fig. 27I ) consists of relatively flat spire whorls and angulated shoulder. Intermediate forms also exist ( Fig. 27B, D, E ). FIG. 24. — Conus ( Kalloconus ) asterousiaensis n. sp. from the Tortonian of Crete (Greece) on apertural, abapertural and apical views, under natural and UV light: paratype AMPG(IV) 2689, Tefeli: the largest specimen with faint colour pattern of dots and dashes; B , holotype AMPG(IV) 2683, Filippi: specimen in abapertural and apical view; C , MNHN.F. A72642 , Crete; D , Paratype MNHN.F. A72644 , Crete; E , MNHN.F. A72647 , Crete. Scale bar: 1 cm. DESCRIPTION OF COLOUR PATTERN Colour pattern on spire whorls consists of two rows of regularly arranged, quadrangular to rectangular dots, near carina and near suture, respectively. Flammulae are visible on some specimens ( Fig. 24B 4 ). The colour pattern on last whorl consists of three layers. The first layer consists of two fluorescent bands, one on the middle of the last whorl and another near the anterior part of the shell ( Fig. 25 ). The second layer consists of multiple, continuous spiral lines, equally distant on individual shells. The lines are not always constant in width. Usually, the thin lines are surrounded by wider lines. The colour of the lines is dim-fluorescent or non-fluorescent. Inside those, there are bright, quadrangular, sharply edged, fluorescent dots alternated with nonfluorescent areas ( Fig. 26 ). The dots are usually as wide as the corresponding line, but sometimes the dots are engulfed by the dim-fluorescent material of the line ( Fig. 26 ). The lines overlap the pattern of the bands ( Fig. 25 ). One specimen shows a third layer of pattern that consists of axially arranged, fluorescent blotches, placed on top of the bands and lines, with dots preserved on top of all other patterns ( Fig. 27C ). FIG. 25. — Elements constituting the colour pattern of Conus ( Kalloconus ) asterousiaensis n.sp. of from the Tortonian of Crete (Greece),with an example of this species’ protoconch,shown under natural light: A , Paratype MNHN.F. A72644 , Crete:Notice the two spiral lines of dots on both sides of the suture,the abrupt transitional pattern of dots in the dim fluorescent,axial lines.Notice the small spiral lines of dots and the fluorescent band on the centre of the shell’s length; B , MNHN.F. A72647 , Crete:this specimen has its multispiral protoconch preserved.Scale bars:A,0.5cm;B,0.1cm. FIG. 26. — Colour pattern of Conus ( Kalloconus ) asterousiaensis n. sp. MNHN.F. A72642 from the Tortonian of Crete (Greece). Notice the large spiral line with the smaller dots at the centre of the line, surrounded by dim fluorescent material. The line is also surrounded by a fluorescent band. Scale bar: 1 cm. TABLE 12. — Comparison of the Tortonian assemblage of Conilithes Swainson, 1840 and Conus ( Kalloconus ) da Motta, 1991 of Crete (Greece) with Neogene (Langhian to Pliocene) assemblages of neighbouring regions (Langhian of Italy ( Sacco 1893a , b; Hall 1966 ), Paratethys ( Harzhauser & Landau 2016 ), Turkey (Erünal-ErentÖz 1958; Landau et al. 2013 ),Tortonian of Italy ( Sacco 1893a , b), Pliocene of Italy ( Sacco 1893a , b). In the column referring to this work (Crete, Greece), the number of examined specimens.

Langhian		Serravallian	Tortonian		Pliocene
Italy	Paratethys	Turkey	Italy	Crete, Greece Italy
Conilithes brezinae ( Hoernes & Auinger, 1879 )	–	•	–	–	49	–
Conilithes herodus n. sp.	–	–	–	–	10	–
Conilithes sp.	–	–	–	–	1	–
Conilithes striatulus ( Brocchi, 1814 )	?	•	–	•	13	•
Conilithes antidiluvianus ( Bruguière, 1792 )	–	•	–	•	1	•
Conus ( Kalloconus ) neumayri Hoernes & Auinger, 1879	–	•	•	–	10	–
Conus ( Kalloconus ) hendricksi ( Harzhauser & Landau, 2016 )	–	•	•	•	39	–
Conus ( Kalloconus ) helladicus n. sp.	–	–	–	–	9	–
Conus ( Kalloconus ) gulemani Erünal-ErentÖz, 1958	–	–	•	–	19	–
Conus ( Kalloconus ) letkesensis ( Harzhauser & Landau, 2016 )	–	•	–	–	1	–
Conus ( Kalloconus ) asterousiaensis n. sp.	–	–	–	*	17	–

REMARKS The Greek specimens present two extreme forms ( Fig. 27A, G, I ), that are connected with intermediate shell forms ( Fig. 27B, D, E ), therefore we regard them as one species. Morphological characters like the subsutural flexure, the smooth spire whorls and the conical straight last whorl characterize this species ( Table 11 ). Despite the morphological variability, the colour pattern remains constant to all shells ( Fig. 27 ). Accordingly, we consider that this species displays a wide variability in spire height and angulation of shoulder, but bears a stable colour pattern variation. The Greek specimens are attributed to the subgenus Kalloconus , because of the lack of the spiral sculpture, a concave conical spire outline, a wide last whorl and broad shoulder. A syntype of Conus raristriatus Bellardi & Michelotti, 1841 , from the Tortonian of Sant’Agata Fossili (BS.038.05.133), was illustrated by Bellardi & Michelotti (1841 : pl. 5, figs 8-9) and later figured by Ferrero Mortara et al. (1984) . The illustration shows a colour pattern of evenly distanced spiral lines, but no spiral rows of dots as on Conus ( Kalloconus ) asterousiaensis n. sp. As such, the syntype suggests that Conus raristriatus and Conus ( Kalloconus ) asterousiaensis n. sp. belong to two different species. However, a Tortonian specimen figured by Davoli (1972 : pl. 8, fig. 1a, b, specimen no. 5545 housed in the Museo Paleontologico dell ‘Universita di Modena’) and named Conus raristriatus displays a shell shape very similar to the morphotypes of Conus ( Kalloconus ) asterousiaensis n. sp. (see Fig. 27B, F, H ). In addition, under natural light the Italian specimen figured by Davoli seems to be displaying similarities of colour pattern (discontinuous spiral lines) with the Greek specimens. For this reason, we consider it conspecific with the Greek material. FIG. 27. — Morphological variation of shoulder and spire height in Conus ( Kalloconus ) asterousiaensis n. sp. from the Tortonian of Crete (Greece). Colour pattern of the specimens seems consistent, despite the fact that there is a wide variety in spire height and angulation of shoulder: A , Paratype AMPG(IV) 2689, Tefeli; B , AMPG(IV) 2688, Filippi; C , MNHN.F. A72645 , Crete; D , AMPG(IV) 2684, Filippi; E , MNHN.F. A72648 , Crete; F , MNHN.F. A72647 , Crete; G , MNHN.F. A72642 , Crete; H , Paratype MNHN.F. A72644 , Crete; I , Holotype AMPG(IV) 2683, Filippi. Scale bar: 1 cm. Conus ( Kalloconus ) asterousiaensis n. sp. could also be compared to Conus ( Lautoconus ) subraristriatus Pereira da Costa, 1866 . They differ morphologically, mainly in the cyrtoconoid, more elongate and higher spire. In terms of colour pattern variations, both species possess the spirally arranged rows of dots and dashes, with fluorescent bands. The difference is that Conus ( Lautoconus ) subraristriatus does not exhibit any pigmentation between the fluorescent bands ( Landau et al. 2013 ; Harzhauser & Landau 2016 ), whereas Conus ( Kalloconus ) asterousiaensis n. sp. possesses colour patterns along the whole length of the last whorl. The colour pattern of Conus ( Kalloconus ) asterousiaensis n. sp. is similar to that of the extant species Conus genuanus Linnaeus, 1758 . This West African species could be related to C. ( K. ) asterousiaensis n. sp. and suggests a Proto-Mediterranean origin of some West African conids.