Late Miocene Conidae (Mollusca: Gastropoda) of Crete (Greece). Part 1: genera Conilithes Swainson, 1840 and Conus (Kalloconus) da Motta, 1991

ABSTRACT Conidae is a diverse family of carnivorous marine gastropods. They rapidly diversified during the Miocene and now they inhabit tropical and subtropical seas. Here we attempt to provide the first inventory of fossil conids from the late Miocene of Crete (Greece). This paper deals with the genera Conilithes and Conus (Kalloconus) da Motta, 1991 and will be followed by papers presenting other genera of the family. Using UV light, we described the residual colour patterns of eleven species, of which three are new: Conilithes herodus n. sp., Conus (Kalloconus) helladicus n. sp and Conus (Kalloconus) asterousiaensis n. sp. One species is in open nomenclature: Conilithes sp.. Six species are first recorded in the late Miocene of Crete: Conilithes brezinae (Hoernes & Auinger, 1879), Conilithes striatulus (Brocchi, 1814), Conus (Kalloconus) neumayri Hoernes & Auinger, 1879, Conus (Kalloconus) hendricksi (Harzhauser & Landau, 2016), Conus (Kalloconus) gulemani Erünal-Erentöz, 1958 and Conus (Kalloconus) letkesensis (Harzhauser & Landau 2016). Conilithes antidiluvianus (Bruguière, 1792) is the only species already recorded by past Greek authors. Firstly, our study reveals that only two species are restricted to the late Miocene of Crete (Conilithes herodus n. sp. and Conus (Kalloconus) helladicus n. sp.). Secondly, we found deep relationships with the conid assemblage from the Langhian of Paratethys (six shared species). This result could be interpreted as a conid fauna, present and widely distributed since the Langhian-Serravallian in both the Paratethys and the eastern Proto-Mediterranean. This fauna disappeared from Paratethys during the Serravallian, but probably persisted in the eastern Proto-Mediterranean, as suggested by the relationships with the Serravallian of Turkey and the Tortonian of Crete (this work). On the other hand, the weak relationships with the late Neogene of Italy might be biased because, for the comparison with Italian faunas, we used works that illustrated Conidae without UV light.


INTRODUCTION
The Conidae snails are a family of marine gastropods with a remarkable biodiversity of more than 900 extant species. Conids are present in a variety of depths in subtropical and tropical seas (e.g., Hendricks 2015;Abalde et al. 2019). They are carnivorous, mainly hunting a specific prey (Kohn 1990). Most species eat polychaete worms, but some lineages eat other gastropods or even fish (Olivera et al. 2015;Safavi-Hemami et al. 2015).
The rich conid fossil record starts from the early Eocene (Duda & Kohn 2005). Kohn (1990) suggested multiple radiation events, but the Neogene is one of the greatest phases of radiation. Until now, Conidae from the Neogene of Greece have sporadically been identified and named as Conus sp., with the few exceptions of Symeonidis (1965), Symeonidis & Konstantinidis (1968), Dermitzakis 1969, Merle et al. (1988, Koskeridou (1997) and Koskeridou et al. (2017). Most of the cone shells were misidentified (Symeonidis 1965;Symeonidis & Konstantinidis 1968), leading to an unclear distribution of the Conidae fauna in the Eastern Mediterranean during the late Miocene. Here, we present the first systematic study of Conidae from the late Miocene of Crete (Greece) using UV light in order to reveal their colour patterns as an aid to their identification. An updated taxonomy of the conids found in the Tortonian of Crete is necessary for an accurate representation of the biodiversity of the family in the late Miocene of Eastern Mediterranean.

GEOLOGICAL BACKGROUND
The island of Crete today is a horst in the forearc of the Aegean region. N-S and E-W extension created normal faults where the footwall is the basement, and the hanging wall Neogene deposits overlie basement rocks of the Upper Nappes. Our study area is in Central Crete, in the Heraklion and Messara Basins (Meulenkamp et al. 1979;Zachariasse et al. 2011) and in two localities from the Ierapetra Basin and Sitia Basin of Eastern Crete (Dermitzakis 1969). Three mountains, Psiloritis Mountains (P), Asterousia Mountains (A) and Dikti Mountains (D), surround the Heraklion and Messara Basins (Fig. 1). Ierapetra graben deposits are overlain on the pre-Neogene series unconformity (Ring et al. 2001). The localities where the fossils were found have been discussed in the past with Symeonidis's work being of great importance for the study of late Miocene mollusc faunas (Symeonidis 1965;Symeonidis & Konstantinidis 1968). 1) Tylissos locality ( Fig. 1[1]) is in the foothills of Psiloritis mountains and according to Delrieu (1990), Tylissos sand facies are covered by marls containing Globorotalia miocenica mediterranea Catalano & Sprovieri 1969. 2) Keramoutsi locality ( Fig. 1[2]) is south of Tylissos, and as well as Tylissos, its sandy sediments are covered by marls containing Globorotalia miocenica mediterranea.

Material
We handpicked samples from the Filippi, Panassos, Apomarma, Tefeli and Makrilia localities. For the other areas we use the historical collections of the National and Kapodistrian University of Athens (NKUA), and the collections of the Muséum national d'Histoire naturelle, Paris, particularly the material coming from the Action spécifique du Muséum project (1989Muséum project ( -1990 on the Neogene of Crete. The specimens were not bleached, as most of them revealed their colour pattern under UV light.

Method
The observation of colour patterns on fossil shells is difficult, as the degradation of the pigments happens quickly after the death of the animal and continues also during fossilisation. A solution to the problem is the use of ultraviolet light (UV), as the colour pattern of the fossil shells becomes visible with the exposure of wavelengths below 365 nm (Miethe & Born  C. et al. 1928;Olsson 1967;Vokes & Vokes 1968;Cate 1972;Hoerle 1976;Hoerle & Vokes 1978;Dockery 1980 Hendricks 2009, 2018; the list is not exhaustive). Concerning the conids, zoologists have long been used the colour patterns to separate extant species, since their shells are conservative in shape and display few shell characters (Marshall et al. 2002;Hendricks 2015). According to Hall (1966), fossil cone shells, that are only observed under natural light and with superficially similar morphologies, have been often misidentified. Consequently, the study of their colour pattern is the best means for their identification at species level (Hendricks 2009(Hendricks , 2015(Hendricks , 2018.

Photographs
We used a CANON EOS-70D with an EFS 15-85 mm image stabilizer ultrasonic lens, using extra magnification × 4 lens. The UV figures were photographed under UV light with wavelength of 365 nm. Also, we occasionally used a LEICA M165 C, with a camera LEICA IC90 E.

Shell terminology
We follow Smith (1930), Röckel et al. (1995, Hendricks (2009), but mainly we used Harzhauser & Landau's (2016) terminology for the subsutural flexure measurements. We follow the 45 angled measurement style of Harzhauser & Landau (2016) in order to compare the Greek specimens with the Paratethys material.

Synonymic and chresonymic lists
We provide the list of the synonyms. Concerning the chresonymic list, we concentrate on Greek references, because, for some species, the list could be very long. This group has received considerable attention recently; the middle Miocene eastern Mediterranean of the Karaman Basin, Turkey was revised by Landau et al. (2013), and the middle Miocene Paratethys by Harzhauser & Landau (2016). In this paper we will not repeat chresonymies and species descriptions given by those authors unless modified or challenged by our findings.   (commonly found in all of the shallow marine Tortonian Formations of Crete

desCription of the Colour pattern
Main colour pattern of this species consists of flammulae on spire whorls (Fig. 3), dots on carina (Fig. 3) and a nonfluorescent band below carina with small to tiny spiral rows of dots (Fig. 3). On the last whorl, this species bears spiral lines of dots usually interspersed with continuous spiral lines, on a non-fluorescent base colour. A non-fluorescent band on the anterior part of last whorl exists in most specimens ( Fig. 2), surrounded by fluorescent bands (Fig. 3). The colour pattern variability of this species results from the ratio of the alternation between continuous spiral lines and other discontinuous lines making small dots (e.g., Fig. 2F, H), as well as the existence of one or two non-fluorescent bands at the last whorl (e.g., Fig. 2C (Landau et al. 2013: 562, pl. 82, fig. 5   shell desCription Small-sized and elongate shells. Protoconch not preserved. Spire with a maximum of ten spire whorls, high, conical with flat sutural ramp in early whorls, slightly concave in later spire whorls. Carina subangulated to angulated, with tubercles visible on early spire whorls, sometimes visible until 8 th spire whorl. Subsutural flexure shallow, strongly curved, strongly to moderately asymmetrical. No spiral grooves below carina. Last whorl elongated, conical. Aperture straight, narrow, widening towards twisted fasciole. Growth lines not prominent, with spiral grooves visible on the anterior part of the shell, towards the anterior part of the last whorl.

desCription of Colour pattern
The colour pattern of the spire whorls consists of thin, axial or irregular fluorescent lines, engulfing angular or irregularly oval, non-fluorescent blotches on carina (Fig. 6). On the body of the shell, two spirally arranged, wide, fluorescent bands exist (Fig. 5), usually disrupted by a non-fluorescent band, with fluorescent blotches or dots. In most cases, the blotches create arrow like patterns (Figs 5G1, 6). Tiny lines of bright fluorescent dots ( Fig. 6) are on the wide fluorescent bands and sometimes on the non-fluorescent base colour, also surrounded by fine thin, continuous, bright fluorescent spiral lines ( Fig. 6).

reMarks
The specimens described herein (Table 2), possess a subangulated to angulated shoulder, revealing a slight morphological variability. The colour pattern on the spire whorls is the most distinguishing character that separates it from the rest of the Conilithes species. The shell of this species is similar to Conilithes brezinae, but none possesses spiral cords below carina, as some Conilithes brezinae specimens do. The colour pattern is different, bearing blotches on spire whorls, two fluorescent bands and one non-fluorescent band in the middle of the last whorl. The similarity in colour pattern on the rest of the shell, bearing lines of dots and continuous spiral lines, suggests a close relation between the species. This species is also morphologically similar to Conilithes sceptophorus (Boettger, 1887), but it differs in its pattern described, consisted of axial zig-zag stripes (Harzhauser & Landau 2016 shell desCription Small-sized, conical shell. Protoconch multispiral (Fig. 7A7). Early spire whorls high, conical, beaded on carina, straight, with a coeloconoid outline (coeloconoid: approaching conical shape but with concave sides). Late spire whorls scalate, straight to convex, with a slightly coeloconoid outline. Carina beaded on the 5-6 early teleoconch whorls only (Fig. 7A).   Maximum diameter at angulated shoulder. Suture channelled. Subsutural flexure shallow, moderately curved, strongly asymmetrical ( Fig. 7A3). Aperture narrow, straight. Last spire whorl straight, conical, narrow. Spiral grooves on the anterior part of last whorl. Fasciole indistinct.

desCription of Colour pattern
Colour pattern consists of thick flammulae on spire whorls. Colour pattern on last whorl consists of fluorescent blotches and closely related, bright fluorescent spiral lines-dashes, that are separated by non-fluorescent elongated dots and nonfluorescent axial blotches (Fig. 8).
reMarks A species differing from Conilithes brezinae and Conilithes herodus n. sp. herein, in the strongly beaded early spire whorls (five or six) and the lower relative height of the spire (RSH) ( shell desCription Small-sized shells with spire whorls of relatively medium height and robust outline. Spire whorls straight to coeloconoid, conical, with scalariform, slightly elevated spire whorls and angulated shoulders. Usual, faint spiral cords on early spire whorls, but no tubercles or beads. Subsutural flexure moderately deep, strongly curved, moderately asymmetrical. Maximum diameter on shoulder. Last whorl straight, conical.  Aperture straight. Fasciole indistinct. Spiral grooves on the anterior part of last whorl.
Colour pattern variation. -The colour pattern consists of fluorescent flammulae on the spire (Fig. 10). The last whorl bears a primary pattern of irregular, fluorescent blotches. The blotches can be axial flammulae that continue from the spire whorls, towards the anterior of the shell, or can be spirally arranged as bands, parallel to a second pattern of fluorescent spiral lines of dots and dashes, along the length of the shell. The spiral rows of dots or dashes start at carina and continue along the length of shell. All patterns are occasionally disrupted by non-fluorescent blotches (Fig. 10). On some shells, a non-fluorescent band exists along the centre of the length of the shell, decorated with fluorescent spiral lines of dots or dashes (Fig. 10

desCription of the Colour pattern
Colour pattern is absent on most of the specimen's surface.
There is a faint pattern of axially arranged, rectangular blotches along the posterior two-thirds of last whorl (Fig. 11A3). reMarks The name of this species has been thoroughly discussed (Janssen et al.

desCription of Colour pattern
The colour pattern consists of one layer displaying very large, polygonal-like, rectangular blotches, restricted axially and spirally (see axial fluorescent and non-fluorescent boundary, Fig. 15). Not all blotches are rectangular. Some have sharp, not straight disruptions, while others fade randomly to the non-fluorescent base colour. On large specimens (see Fig. 15), some blotches tend to faintly connect with each other with faintly fluorescent areas between blotches (see unclear spiral interactions, Fig. 15). The pattern is continuous from the anterior part of the last whorl to the spire (Fig. 13A6). Blotches might be narrow, separated by two non-fluorescent spiral bands, thus creating dash-like rows of blotches.
reMarks This species is not common in Crete (Table 12), but is very easily recognizable under UV light. The colour pattern of large rectangular blotches is characteristic of the species. Moreover, the interactions between the blotches and the dash-like patterns are also characters of this species (Figs 13, 15; see also Harzhauser & Landau 2016: fig. 11E1, F1). The Greek specimens differ morphologically from the Paratethyan ones in the strongly asymmetrical subsutural flexure (Table 6; moderately asymmetrical on Harzhauser & Landau 2016), but we consider that this difference could result from a local variation. Landau et al. (2013), in our opinion, misjudged the more extreme Conus (Kalloconus) neumayri pattern (e.g., Fig. 14). They consider this extreme pattern as that of Conus (Monteiroconus) daciae from the Karaman Basin, Turkey (see Landau et al. 2013: pl. 81, fig. 6a, b). The colour pattern described therein is identical with the pattern of Conus (Kalloconus) neumayri. Unfortunately, their assumption was not fixed in Harzhauser & Landau (2016), since they assumed that Conus (Kalloconus) neumayri shows no signs

desCription of Colour pattern
The colour pattern consists of one layer of fluorescent, evenly arranged rows of dots. The dots are evenly spaced, evenly sized, differing in shape. Some dots have an oval shape; others are more rectangular to parallelogram, while a few are arrow- like shaped (Fig. 17). The axial distance between individual rows does not change with the individual's growth. Newly developed spiral lines of dots are added to fill the gaps, seen as faded, tiny dots between two rows (Fig. 17). As a result, large specimens tend to have numerous rows of dots, while smaller specimens have less rows. The largest specimen has over 22 rows (abapical rows are not clearly visible), while younger have less than 15. On the spire whorls, there is one spiral row of dots, with most of those partly covered by the suture of the succeeding whorls (Fig. 16). for Conus berghausi because of the presence of spiral cords on spire whorls. The study material (Table 7) fits the Conus (Kalloconus) hendricksi shell morphology, on the constrained, defined shoulder and the smooth, coeloconoid early spire whorls (Fig. 16). In the study material, the pointed, early spire whorls are absent on adult specimens, possibly because of the destruction and erosion, of their early, pointed whorls (Fig. 16A, B). This might cause confusion and misleading results in PCA analysis (see Harzhauser & Landau 2016). Thus, we refrain from using this method on the Greek material. Furthermore, Conus (Kalloconus) hendricksi has a consistent colour pattern (Harzhauser & Landau 2016). Harzhauser & Landau (2016) described 13-16 spiral lines of dots on the last whorl (Conus (Kalloconus) hendricksi paratype), with some of them bearing smaller dots. On the Greek specimens, we observe more lines of dots (22 rows visible), and we consider that this small difference probably results from a geographical variation of the character. Finally, the spiral dots on the spire whorls of our Greek specimens, match with Conus (Kalloconus) hendricksi. For these reasons we attribute them to this species.
The specimen of Conus (Monteiroconus) berghausi from the Karaman Basin (Turkey) figured by Landau et al. (2013)   displays a colour pattern which is identical to the study material. In their figure showing the colour pattern of the species (Landau et al. 2013: pl. 81, fig. 1), the last row of dots near the suture is near or above the shoulder. This means that this specimen has spiral rows of dots on its sutural ramp, the characteristic pattern of Conus (Kalloconus) hendricksi and not flammulae, a pattern character described on Conus shell desCription Medium-sized, robust shells, with relatively low spired whorls. Early spire whorls coeloconoid. Last spire whorls, smooth, straight to concave, creating a low conical to flat outline. Suture impressed. Subsutural flexure shallow, weakly curved, moderately asymmetrical. Shoulder rounded, protruded, creating a bulky outline. Maximum diameter below shoulder. Last whorl straight. Aperture moderate, narrow near suture, straight. Apertural canal wide, fasciole twisted, demarcated from base and inner lip. There are two extreme forms. Form 1 consists of robust forms which are relatively wider in comparison to form 2 and have low angled spire whorls. Form 2 consists of relatively elongated forms with flat spire whorl. Intermediate forms also exist.

desCription of Colour pattern
The colour pattern consists of one layer of short and long, fluorescent, spiral dashes, arranged in evenly spaced spiral rows. The spire whorls display wide, fluorescent flammulae, with irregular boundaries on a non-fluorescent base colour. The flammulae do not connect with the colour pattern of the last whorl.
reMarks This species shows some variations in the relative diameter of its spire whorls (Table 8). The difference between the elongated and robust forms is not very variable. However intermediate forms (Fig. 18B, C) between both forms (Fig. 18A, D) point towards the existence of a single species. Conus (Kalloconus) hungaricus specimens sensu Landau et al. (2013: pl. 37, figs 9, 10, pl. 38, fig. 1) from the Karaman Basin (Turkey) are more likely to be Conus (Kalloconus) helladicus n. sp., because of their flat spire whorls and their identical colour pattern.

desCription of Colour pattern
The colour pattern consists of one layer of a series of closely related, spiral rows of dashes (Fig. 20B), disrupted randomly by non-fluorescent dots or small dashes. The non-fluorescent dots are slightly wider than the fluorescent spiral rows. Sometimes the dots are axially aligned, creating a synchronous, vertical disruption of the spiral rows. The dots on the spiral rows are not constant in numbers or distances and can be multiple or few. This results in a variety of colour patterns, with shells having mostly spiral rows of elongated dashes with very few interruptions (Fig. 20B), to patterns with multiple disruptions, resembling series of short fluorescent dashes (Fig. 20D).
The colour pattern of this species is similar to species like Conus (Kalloconus) hungaricus Hoernes & Auinger, 1879 and Conus (Kalloconus) tietzei Hoernes & Auinger, 1879, but both species differ from Conus (Kalloconus) gulemani by their shell morphology (Fig. 21). Conus (Kalloconus) hungaricus has club shaped shells, wider relative diameter of the last whorl and conical spire whorls. Conus (Kalloconus) tietzei has a more angulated shoulder and straight last whorl, which is slightly   spire whorls is partially destroyed on this specimen, but most likely consists of fluorescent flammulae (Fig. 22).
reMarks This species has a low spire and a broad, conical last whorl, with smooth shoulder and a flat-sided last whorl (  fig. 10F-H). These bands are not visible on the Cretan specimen. The presence of this species in the Tortonian Eastern Protomediterranean, is an addition to the important cohort of taxa common to both the Middle Miocene of the Paratethys and eastern Proto-Mediterranean. The typical Paratethyan Langhian assemblages (Harzhauser & Landau 2016) were found in the Proto-Mediterranean in the Serravallian of Turkey (Landau et al. 2013). This species is an example of the persistence of some taxa into the late Miocene of Eastern Proto-Mediterranean.

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. 24B4). 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) (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).

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.  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   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. 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.

CONCLUSION
The study using UV light of the conid genera Conilithes and Conus (Kalloconus) from the late Miocene of Crete (Greece) reveals a high species diversity from the tropical environment of the Proto-Mediterranean. The collection of the NKUA actually contains a much more diverse variety of fossil Conidae than the one described by past research collectors, because they could not differentiate species in natural light. In this work, we recognize eleven species, among which one is left in open nomenclature (Conilithes sp.), three are new (Conilithes herodus n. sp., Conus (Kalloconus) helladicus n. sp. and Conus (Kalloconus) asterousiaensis) and six are recorded for the first time in the late Miocene of Crete (Table 12). We compare this assemblage with those of the Miocene and Pliocene neighbouring regions (Italy, Turkey and Paratethys; Table 12). First, only two species (Conilithes herodus n. sp., Conus (Kalloconus) helladicus n. sp.) are endemic from the late Miocene of Crete. Secondly, we found strong relationships with the Langhian of Paratethys (six shared species), whereas three species are shared with the Serravallian of Turkey (Karaman Basin) and only four with the Tortonian of Italy. This result could be interpreted as a fauna, present and widely distributed since the Langhian-Serravallian in both the Paratethys and the eastern Proto-Mediterranean (Landau et al. 2013). This fauna disappeared from the Paratethys during the Serravallian (= Sarmatian), because of strong environmental changes due to the disconnection from the Proto-Mediterranean (Landau et al. 2013), but probably persisted in the eastern Proto-Mediterranean, as suggested by the faunas of the Serravallian of Turkey and the Tortonian of Crete (this work). However, the weak relation of the Cretan fauna with the late Neogene of Italy could be biased, because, for the comparisons with Italian faunas, we use works that figured Conidae in natural light, as no recent works used UV light to study conids. Consequently, this disparity of information between the Paratethys and Italy could be another reason why we found more affinities with the Langhian of Paratethys than with the Tortonian of Italy.