Late Miocene Conidae (Mollusca: Gastropoda) of Crete (Greece). Part

. Late Miocene Conidae of Crete (Greece) have recently been evaluated for the genera Conilithes and Conus ( Kalloconus ). We continue this ﬁ rst inventory of the Miocene Conidae from Crete by discussing the genera Conus ( Lautoconus ), Conus ( Stephanoconus ), Conus ( Plagioconus ) and two species, not attributed to a subgenus of Conus . With the use of UV light, we recognized 17 species, of which ﬁ ve are new: Conus ( Lautoconus ) ictini sp. nov., Conus ( Lautoconus ) lauriatragei sp. nov., Conus ( Lautoconus ) damianakisi sp. nov., Conus ( Stephanoconus ) moissettei sp. nov. and Conus davolii sp. nov. Six species are ﬁ rst reported in the Late Miocene of Greece: Conus ( Lautoconus ) cf. baldichieri Borson, 1820, Con us ( Stephanoconus ) cf. taurinensis Bellardi & Michelotti, 1841, Conus fuscocingulatus Hörnes 1851, Conus ( Plagioconus ) elatus Michelotti, 1847 and Conus ( Plagioconus ) aquensis d’Orbigny, 1852. Six species are left in open nomenclature. on last whorl. Aperture straight, relatively narrow. Fasciole carved by growth lines. Last whorl straight conical.


Introduction
Conida e Fleming, 1822 snails are carnivorous gastropods that prey mainly on bristle worms (Polychaeta), while some species hunt other gastropods or fi sh Safavi-Hemami et al. 2015). Their rich biodiversity, reaching more than 900 extant species, inhabits a wide range of depths in subtropical and tropical seas (e.g., Kohn 2 014; Monnier et al. 2018;Abalde et al. 2019). Their fossil record starts from the Early Eocene (Duda & Kohn 2005) and it has been suggested that they radiated multiple times in the past (Kohn 1990), with the greatest radiation event happening in the Neogene. There is little information on the Miocene Conidae of Greece, with limited references (Symeonidis 1965;Symeonidis & Kostantinidis 1968;Dermitzakis 1969;Merle et al. 1988;Koskeridou 1997;Koskeridou et al. 2017;Thivaiou et al. 2019) where they have often been misidentifi ed. The unclear diversity of the Conidae of Greece has been tackled by Psarras et al. (2021), who revised the genera Conilithes Swainson, 1840 and Conus (Kalloconus) da Motta, 1991, using UV light to reveal their colour patterns. The di versity of colour patterns is helpful to distinguish many living species and is important for the identifi cation of fossil conids (e.g., Hendricks 2015;Psarras et al. 2021 for the Miocene of Crete). In this work, we continue the systematic revision of the Tortonian Conidae from Crete by showcasing the subgenera Conus (Lautoconus) Monterosato, 1923, Conus (Stephanoconus) Mörch, 1852 and Conus (Plagioconus) Tucker & Tenorio, 2009.

Geological background
Crete is located in the southern part of the Aegean, interpreted as a horst structure in the forearc of the Hellenic plate (Meule nkamp et al. 1979;Zachariasse et al. 2011). Due to normal faults in N-S and E-W directions formed, the Upper nappes are acting as the basement's footwall, while the Neogene deposits are acting as the hanging wall. The specimens were collected from the Heraklion and Messara Basins in Central Crete, and the Ierapetra Basin and Sitia Graben in Eastern Crete. The Heraklion and Messara Basins are surrounded by the Psiloritis Mountains to the North-West, the Asterousia Mountain range to the South and Dikti Mountains to the East. The two basins are divided by the E-W striking Central Heraklion Normal Fault (CHNF) that was active between 9.7 and 7.36 Ma (Zachariasse et al. 2011). In the Ierapetra Basin, the Neogene deposits are deposited unconformably above the pre-Neogene series (Ring et al. 2001). The Apostoli Basin was shallow with lacustrine, brackish and shallow marine deposits during the Tortonian (Drinia 1998). The localities mentioned in Psarras et al. (2021) are also discussed herein (Fig. 1).

Material
We handpicked samples from four localities: Filippi, Panassos, Apomarma and Tefeli. For the other localities, we studied the historical collections of the National and Kapodistrian University of Athens (NKUA), and the Muséum national d'histoire naturelle, Paris, in particular the specimens collected during the 'Action spécifi que du Muséum project ' (1989-1990) on the Neogene of Crete. Most specimens from Parti ra and Tefeli were collected by one of us (E.K.). If the colour patterns of the fossil specimens are not clearly visible under UV light, the specimens can be bleached using sodium hypochlorite, so their patterns could be revealed (e.g., Caze et al. 2015). In this case, it was not necessary to bleach the specimens, as most of them already display a colour pattern under UV light.

Methods
Conidae from the Tortonian of Crete can be characterised by using a combination of morphological and colour pattern characteristics visible under UV light, as was observed on other well-preserved fossil gastropods (e.g., in older works: Ol sson 1967; Vokes & Vokes 1968;Krueger 1974;Hoerle 1976, or recent works: Merle et al. 2008Caze 2010;Caze et al. 2011aCaze et al. , 2011bCaze et al. , 2015Landau et al. 2013;Hendricks 2015Hendricks , 2018Harzhauser & Landau 2016). When gastropods die, the pigments on the shell surface degrade, but don't totally disappear. By exposing the shells under ultraviolet light, their pigments become fl uorescent, although their precise nature is most likely still unknown (Affenzeller et al. 2019). The contrast between fl uorescent and non-fl uorescent areas on the shell enhances the revelation of a residual colour pattern, almost similar to that of the shell surface, when the animal was alive. This method is very useful in the identifi cation of the Cretan conids. Without disregarding the morphology of the shell, which is the primary characteristic for identifi cation, the method of colour pattern identifi cation is a useful addition to our 'researching artillery' that practically helps disentangle clusters of species which are conservative in shape (Marshall et al. 2002), but possess different colour pattern variations (Gong et al. 2012;Hendricks 2015). For the identifi cation of species, we fi rst studied specimens and subsequently grouped them into groups according to their morphology and colour pattern variations. Then, we did statistical analysis using Principal Component Analysis (PCA) graphs to verify our interpretations. By imputing measurements for each specimen, the program PAST computes the overall outline of each shell and groups specimens into a two-dimensional graph. The interpretation of such clusters usually displays species' morphological variation. The input of the data, here used, consists of different ratios of measurements concerning the length, diameter and the apertural height of each shell. Despite some ratios having the same underlying measurements, the resulting graphs successfully display the differences between the group.

Photographs
We photographed with a CANON EOS-70D with an EFS 15-85mm image stabilizer ultrasonic lens, adding when necessary extra magnifi cation 4 × lens. The fi gures were taken using decreased exposure levels (-2), in order to better capture the UV colours. Multiple photographs were taken at different focus points, which were then combined and blended into Photoshop. The UV photographs were taken under UV light with wavelength of 365 nm. The UV lights mostly used are two fl ashlights Convoy S2+, placed diagonally on both sides of the fi gured specimen. On both fl ashlights the original glass was replaced with a ZWB2 fi lter, which blocks any natural light emitted from the fl ashlight. The fi gures from MNHN wer e taken using two ATLAS MMT GmbH Fluotest Forte UV lamps, placed diagonally on both sides of the fi gured specimen. Also, we occasionally photographed specimens under a LEICA M165 C stereoscope, with a camera LEICA IC90 E.

Shell terminology
We follow the terminolog y of Smith (1930), Röckel et al. (1995) and Hendricks (2009), for the description of the shell's dimensions. The shell lengths are considered: large (> 80 mm), moderately large (55-80 mm), medium-sized (35-55 mm), moderately small (25-35 mm), small (15-25 mm) and very small (< 15 mm). For the subsutural fl exure measurements, we used Harzhauser & Landau (2016), following their 45°-angled measurement style in order to compare the Cretan material with the Paratethys material. The identifi cation of species has been done by using as many characteristics as possible, therefore all measurements are equally important. The subsutural fl exure, even though it is variable at the intraspecifi c level and even differs in individuals, is an equally important tool for the description of a species. For the systematic arrangement we follow Puillandre et al. (2014Puillandre et al. ( , 2015 and refer to Lautoconus, Stephanoconus and Plagioconus as subgenera, as in Psarras et al. (2021). The colour patterns which show two to four different colours are called, according to Meinhardt (1998), "levels of pigmentation".

Measurements
We follow the terminology of Smith (1930), Röckel et al. (1995) and Hendricks (2009), and Harzhauser & Landau (2016). The measurements were taken from individual shells for comparison with other fossil Conidae from Europe. For the ratios we follow Harzhauser & Landau (2016), which we use for the PCA graphs.

AH
= apertural height AL = apertural length HMD = height of maximum diameter LW = length-width ratio LWA = last whorl angle MD = maximum diameter PMD = position of maximum diameter (HMD/AH) PV = position of the vertex RD = relative diameter (MD/AH) RSH = relative height of the spire {(SL-AH)/SL} SA = spire angle SL = shell length SSFD = subsutural fl exure depth SSFd = subsutural fl exure relative depth The measurements of the studied specimens can be found in the Supplementary fi le 1.

Diagnosis
Moderately small to moderately large, turnip-shaped species. Medium-wide to wide shells and convex sides. Spire low to moderately high. Paucispiral protoconch. Early whorls ornamented with cords, which might persist on later spire whorls or appear as spiral threads. Subsutural fl exure from very shallow to deep, usually moderately curved, moderately asymmetrical. colour pattern variable, displaying blotches, spiral rows of dots, dashes, and tents in multiple combinations.

Remarks
Puillandre et al. (2014,2015) treated Lautoconus at the subgenus level. In this work we show two species that are large-sized (Conus (Lautoconus) damianakisi sp. nov. and Conus (Lautoconus) lauriatragei sp. nov.). The species of the subgenus display an intraspecifi c morphological variability, as it is here shown for more than one species. The colour patterns are also variable, displaying patterns such as blotches, spiral rows of dots, dashes and tents in multiple combinations (see Monnier et al. 2018). We follow Harzhauser & Landau (2016) in excluding from this subgenus the species with tuberculate spire whorls. The oldest occurrence in Greece is reported from the early Miocene (Thivaiou et al. 2019) by an incomplete specimen (AMPG(IV) 2473). However, the attribution to Lautoconus seems doubtful and this specimen could possibly b elong to Conus (Eoconus) Tucker & Tenorio, 2009 (see more in Tracey et al. 2017).

Shell description
Small-medium-sized (SL: 39.57 mm) and elongated shell, with straight to cyrtoconoid spire outline. Spire convex, conical of variable height, ranging from nearly fl at to low conical. Early spire whorls straight, striated. Late spire whorls convex, with impressed suture. Subsutural fl exure very shallow, weakly curved, moderately asymmetrical ( Fig. 40A). Shoulder rounded, with maximum diameter varying in position, from near shoulder to one fourth posterior part of last whorl. Last whorl slightly infl ated. Aperture straight to slightly curved, widening considerably towards fasciole. Siphonal canal wide, straight. Fasciole twisted, slightly infl ated in larger specimens. Spiral cords present along the anterior part of last whorl.

Description of colour pattern
The colour pattern on spire whorls consists of periodical, irregularly shaped dashes, sometimes parallel to the direction of shell growth. The colour pattern on the rest of the shell consists of spiral rows of fl uorescent dashes, demarcated by non-fl uorescent areas. Faintly fl uorescent blotches appearing sporadically in some shells, surrounding the dashes and connecting axially arranged spiral dashes. Axially arranged spiral dashes tending to form vertical, diagonal, or zigzag axial formations (Fig. 3). Occasionally, a hint of a non-fl uorescent band on middle whorl height, engulfi ng a spiral line of short dashes (seen clearly on Fig. 2E).

Remarks
The Cretan specimens differ from the other species of Conus (Lautoconus) herein, in the relatively slender morphology (Table 1), the low angled spire and the exquisite colour pattern of delicate spiral rows of dashes. We argue that the morphological variability recorded in the studied material is likely due to the different overall shape during development, with larger specimens having a more robust outline. Small specimens which are more elongated, could be interpreted as Lautoconus pyruloides (Sacco, 1893a) (Ferrero Mortara et al. 1984 fi g. 9, pl. 16 fi g. 7), but past researchers assigned them to Conus (Lautoconus) eschewegi (Hall 1966;Davoli 1972). In Greece, specimens studied by Symeonidis & Konstantinidis (1968) (2016) mentioned that Pereira da Costa had a "much too broad species concept". We disagree with that statement as well, because in the Cretan specimens there is also a spectrum of variability in morphology, with smooth to slightly angulated shoulder, fl at to infl ated spire whorls and a conical to concave spire. This variation has also been shown in specimens of Davoli (1972: pl. 6 fi gs 1-16), indicating a wide morphological variation for this species.

Stratigraphic range
Burdigalian of Italy (Hall 1966

Diagnosis
A species with smooth, low angled, conical spire whorls and bricked-wall-like colour pattern.

Etymology
Name taken after Ictinus (Latin translation of the ancient Greek Ἰκτῖνος), one of the two architects of the Parthenon.

Shell description
Moderately small to medium-sized (H. max: 42.68 mm), oval to olive-like shells, with smooth spire whorls. Spire conical, smooth, slightly elevated in some specimens. Early spire whorls coeloconoid in outline, individual sutural ramps straight to slightly convex. Late spire whorls convex, with straight outline. Suture incised. Subsutural fl exure shallow, moderately curved, moderately asymmetrical ( Fig. 40B). Shoulder rounded to slightly angulated. Maximum diameter right below shoulder. Last whorl elongated, smooth. Spiral cords on anterior part of last whorl. Fasciole twisted distinct, swollen.

Description of colour pattern
The residual colour of the last whorls consists of three levels of colouration. The fi rst vivid colouration is made up of horizontally arranged, densely packed, evenly spaced, continuous spiral lines (Fig. 5) and vertical to diagonal, narrow, short lines that connect the spiral lines, creating a bricked-wall-like pattern (Fig. 5). This pattern is faded in many parts of the shell (Fig. 4B). Under natural light, though, the colour pattern is visible even without the help of UV light on the holotype as parallel, closely arranged lines (Fig. 4A1, A3). A second level of colour consists of fl uorescent blotches on the last whorl and irregular fl ammulae on the spire whorls. The fl uorescent blotches are often fi lling the rectangular areas between the spiral lines of the fi rst level of colour ( Fig. 4C-D). The third level consists of a non-fl uorescent pattern bearing irregular blotches that overlaps the other patterns (Fig. 5).

Remarks
Morphologically (Table 2), this species is very similar to Conus (Lautoconus) pelagicus Brocchi, 1814. It differs slightly in the smoother spire whorls and a more infl ated appearance (Fig. 4). The occurrence of this morphotype, at multiple localities and ages (Serravallian, Turkey (Landau et al. 2013) and Tortonian, Greece (this work)), is evidence for the existence of a separate species.

Description of colour pattern
The colour pattern consists of closely related, delicate, elongated, wavy spiral rows of dashes. On the spires, the pattern consists of very thin axial fl ammulae, closely related to each other. The fl ammulae are vaguely connected with a spiral band, at the center of the sutural ramp.

Remarks
This species is very similar in morphology (  Table 3. Shell measurements of one broken specimen of Conus (Lautoconus) sp. 1 (MNHN.F.A82982) from the Tortonian of Crete (Greece).

Shell description
Medium-sized shells (51.92 mm), with medium to low conical, straight to slightly coeloconoid spire. Early spire whorls faintly striate, highly pointed. Later spire whorls smooth, straight to slightly coeloconoid. Faint spiral grooves on spire whorls, diminishing on late spire whorls. Suture channelled. Subsutural fl exure shallow, moderately curved, moderately asymmetrical (Fig. 40D). Shoulder rounded, with an angulation at suture height. Maximum diameter right below shoulder. Last whorl conical, with variable elongation of the last whorl, not con stricted. Spiral grooves present on anterior third of last whorl. Aperture moderate, widening anteriorly. Fasciole indistinct, slightly twisted.

Description of colour pattern
The colour pattern c onsists of two levels of colouration. The fi rst consists of ye llow axial stripes on a non-fl uorescent base. Those stripes are either continuous from the shoulder, where a non-fl uorescent disruption exists (Fig. 8), until the anterior of the shell, or are not continuous, showing patterns of axial disruptions, convergences with other stripes, and divisions of individual stripes into more (Fig. 8A). The second level consists of two axial bands; the fi rst one is placed just above the middle of the shell and the second one is placed on the anterior part of the shell, along with the last spiral groove. The bands are not always continuous or visible (Fig. 8B). In some shells, they resemble closely arranged, thin spiral dashes ( Fig. 8E, G-H), some of them axially arranged (Fig. 8H).

Remarks
This species is the most common Conus (Lautoconus) in the Tortonian of Crete. The study of more than 100 specimens (Table 4) consistently shows the recurrence of the morphology and colour pattern of this species (Fig. 8). From the Serravallian of Turkey, Erünal-Erentöz (1958: 121, pl. 20 fi gs 4-5) described Conus (Chelyconus) pyrula var. mucronata, but the binome Conus mucronatus is occupied by a recent species, described by Reeve (1843). The Turkish material displays a morphology similar to the Cretan specimens, with pointed early spire whorls, low conical late whorls and an angulated shoulder. As no colour pattern has been studied for that material, we cannot consider with certainty that the Cretan and the Turkish specimens belong to same species. The type specimen of the Pliocene Conus bitorosus var. elatoastensis Sacco, 1893(Sacco 1893b, illustrated by Hall (1966: pl. 23 fi g. 24), is very similar to Conus (Lautoconus) sp. 2, but we have not observed its colour pattern.
The shell morphology of the Pliocene Conus (Lautoconus) ponderosus Brocchi, 1814 resembles that of Conus (Lautoconus) sp. 2, but differs by lacking the early, pointed spire whorls, by the occurrence of spiral grooves on last whorl and by the frequent angulation on shoulder. Its colour pattern seems not preserved (Harzhauser & Landau 2016; Annalaura Pistarino pers. comm.). One syntype of Conus (Lautoconus) conoponderosus Sacco, 1893b (MRSN BS.038.05.082, Tortonian of Colli Tortonesi) was fi gured in Davoli (1972: pl. 4 fi g. 19). It bears a similar morphology, but displays a lower spire and a stout last whorl differing from most of the specimens studied here. Its colour pattern was not observed. Despite the discussed differences in shell morphology, Conus ponderosus and Conus conoponderosus might be different morphotypes of a same species. Since we observed similar morphotypes from Crete, we cannot disregard the possibility of them being conspecifi c. Thus, we refrain from attributing the Cretan material to Conus (Lautoconus) ponderosus, Conus (Lautoconus) conoponderosus or Conus elatoastensis. Further examination needs to be done on the Italian types by using UV light in order to clarify whether the colour pattern allows distinguishing these species.

Colour pattern variation
Three levels of colouration are present in this species. The fi rst one consists of two fl uorescent bands, which are on the middle and anterior parts of the shell. The second one is a pattern of spirally arranged alterations of non-fl uorescent and fl uorescent dots. The third level is displayed between the spiral alterations of dots. It consists of a series of fl uorescent, arrow-like blotches-tents, pointing in the opposite direction of the shell growth, alternated by non-fl uorescent areas. The non-fl uorescent areas might have one or two dots between the fl uorescent arrows. Along with the second level of fl uorescent dots, irregular fl uorescent fl ammulae appear on the spire whorls.

Remarks
The colour pattern of the species is unique in the studied material. The relatively rounded spire whorls and the lack of tubercles on early spire whorls allow us to categorize it as an elongated Conus (Lautoconus).
Caze (2010) 'Orbigny, 1852), in the higher spire and the more angulated shoulder. Finally, this species is not Conus (Lautoconus) ponderosus Brocchi, 1814 as it has a slightly higher spire and more elongate last whorl.

Description of colour pattern
The colour pattern consists of three levels of colouration. The fi rst one consists of continuous grouped spiral lines of dim-fl uorescent colour (Fig. 12, dim fl uorescent lines). The second one consists of continuous spiral lines of bright-fl uorescent colour (Fig. 12, bright green coloured lines). Both patterns of lines are slightly wavy, and it is possible for the second one to overprint the fi rst. The third level of colouration is a series of thin, wavy, fl uorescent and axial lines (Fig. 12, axial lines). The lines are not always continuous, but are more prominent on the spire whorls. Some of the lines are linked one to another, turning into a pattern of axially entangled lines. The result of the three patterns reminds of vehicle tire tracks (Fig. 11A2).

Remarks
Ferr ero Mortara et al. (1984: pl. 16 fi g. 8) fi gured the 'holotype' of Conus baldchieri (in fact they made a fi xation of a lectotype by inference of holotype, ICZN 1999: Art. 74.6)) regarded by Sacco (1893a) as Lithoconus mercatii var. baldichieri. The Cretan specimen attributed to Conus (Lautoconus) cf. baldichieri is similar to Conus baldichieri Borson, 1820 from the Pliocene of Italy, but differs in the less striated spire whorls, the less infl ated shell, the more elongate last whorl and its shorter siphonal canal.

Shell description
Large-sized shell (SL max.: l83.84 mm), with hemispherical-shaped spire and smooth shoulder. Early spire whorls convex, smooth, resulting in fl at to convex outline. Later spire whorls strongly convex, elevated. Suture channelled, irregular. Subsutural fl exure very deep, moderately curved, strongly asymmetrical (Fig. 40G). Shoulder smooth, with maximum diameter at shoulder. Aperture narrow, curved. Last whorl conical, straight to concave, narrowing slightly at centre of last whorl. Deeply carved growth lines on last whorl. Aperture not straight, wavy. Fasciole very small, to indistinct.

Description of colour pattern
Shell surface partly damaged (Fig. 13). The visible colour pattern consists of wide, axial fl ammulae on the spire whorls, that continue axially as ribbons on the body of the shell. The ribbons are connected with irregular fl uorescent blotches (Fig. 14).

Remarks
This species is quite unusual in morphology (Table 7), being very large with a semi-circular spire outline species of Conus (Laut oconus) with a similar morphology occur today, such as Conus (Lautoconus) regonae Rolán & Trovão in Rolán, 1990 (see Tenorio et al. 2020: fi g. 6d). Therefore, we are confi dent in assigning it to this subgenus. Other than the Cretan specimen, another specimen called Conus (Lautoconus) sp. (HNHM INV 2013.291;Kovács & Vicián 2013) has also been found in the Paratethys from the Pannonian Basin at Letkés, Hungary, and for us it could belong to the same species. Its colour pattern is not comparable to any of the Conidae studied herein. Despite not having at least

Shell description
Large shells (SL max.: 80.02 mm), with smooth, conical spire whorls and spiral cords. Spire whorls straight to slightly convex, with slightly concave outline in early spire whorls, straight to slightly convex in later spire whorls, with smooth to slightly elevated interactions of spire whorls, creating smooth, low conical to convex outline. Multiple spiral cords on top of spire whorls, with three to four on early spire whorls, up to ten on late spire whorls. Shoulder rounded. Maximum diameter below shoulder. Suture, channelled, not straight. Subsutural fl exure moderately deep to shallow, moderately curved, strongly to moderately asymmetrical (Fig. 40H). Last spire whorl conical, straight. Aperture narrow, fasciole small, slightly twisted. Spiral grooves near anterior part of last whorl.

Colour pattern variation
The colour pattern on the spire whorls consists of thin fl ammulae, frequently very closely related to united, resulting in wide, fl uorescent and comma shaped fl ammulae (Fig. 15, see apical views). The colour pattern of the last whorls consists of two levels of pigmentation. The fi rst one consists of wide, dim-fl uorescent spiral bands, interrupted by non-fl uorescent, thin spiral, continuous bands or lines. The second level consists of very closely related, fl uorescent dashes to continuous lines. The dashes tend to be on an axial synchronization with other spiral dashes-lines. The width of the spiral bands varies, as well as the number of interruptions of the spiral lines. Also, the length of the dashes varies as well, from almost dot-shaped dashes to continuous spiral lines (Fig. 16).

Remarks
The characters of t he spire whorls and the colour pattern variation differentiate this species from the rest of the studied material. The specimens are similar in morphology (Table 8) to Conus pseudo-textilis Grateloup,195 8. The studied species differs from the variety of Erünal-Erentöz in the slightly concave sutural ramps. This difference is crucial, because the concave sutural ramps is shared between the specimens of Erünal-Erentöz and members of Monteiroconus, but not the members of Lautoconus.

Shell description
Medium-sized shell (SL max.: 29.43 mm). Early spire whorls conical to convex, of low to medium height. Later spire whorls conical in outline, with convex, smooth sutural ramp. Subsutural fl exure moderately deep, strongly curved, moderately asymmetrical (Fig. 40I). Shoulder smooth at early spire whorls, slightly angulated at later spire whorls. Maximum diameter below shoulder. Last whorl conical, slightly widened. Aperture narrow, straight, widening towards slightly twisted fasciole. Shell smooth, except for two faded spiral cords near fasciole.

Colour pattern variation
The colour pattern on the spire whorls consists of irregular blotches. The colour pattern on the last whorl consists of two levels of colouration. The fi rst one consists of fl uorescent blotches. The second level consists of spirally arranged irregular rows of fl uorescent dashes, interrupted by non-fl uorescent tents.
The tents can be either as small as dots, or small non-fl uorescent dashes, or can be axially wide enough to unite with other tents and create an axial non-fl uorescent area. The resulting pattern is variable and depends on the number of tents, as well as on the size of those tents. The Cretan specimen shows an axial unifi cation of the tents, while the Turkish specimen displays a spiral unifi cation of the tents, in the middle part of the last whorl.
We noticed that the specimen from Apomarma (MNHN.F.A83015) had different grades of erosion, with non-fl uorescent areas being more resistant to erosion, than those being previously with colour patterns (see Fig. 17A4 in comparison to the right side of Fig. 17A3, where the shell is not so eroded). This might indicate that the substance enclosed in the shell structure, which is responsible for the pigments, is more prone to erosion than the non-pigmented areas.

Remarks
This species is placed in Lautoconus, as it has smooth early spire whorls and a convex sutural ramp. The Pliocene Turkish specimen identifi ed by Caze (2010) as Conus (Chelyconus) ?pyrula Brocchi, 1814 and the Cretan specimen of Conus (Lautoconus) sp. 4 share smooth and convex spire whorls with a slightly angulated shoulder (Fig. 17) and have a similar colour pattern. Thus, they very likely belong to the same species. The typical Pliocene Conus pyrula bears a colour pattern of axially arranged stripes. Conus (Lautoconus) sp. 4 differs from the Paratethyan species of Conus (Lautoconus) (Harzhauser & Landau 2016) in the presence of tents in its colour pattern. Conus (Lautoconus) sp. 5 (see below) has a similar colour pattern of tents, but it differs morphologically from this species in the smooth outline of the spire whorls and the olive-like morphological outline. As such, we consider these specimens as two separate species. Unfortunately, in lacking more specimens with similar morphological characteristics and colour pattern, we do not name this species. Conus (Lautoconus) sp. 4 has a very common shell outline and is not easily distinguishable from other species without the help of UV light. The colour pattern is also reminiscent of extant species of Conus (Lautoconus), such as the West African species Conus (Lautoconus) saragasae Rolán, 1986(Tenorio et al. 2020.

Description of colour pattern
The colour pattern on the spire whorls consists of fl uorescent irregular blotches and, inside those, randomly positioned, non-fl uorescent tents. The tents continue until shoulder height. The colour pattern on the last whorl consists of two levels of pigmentation. The fi rst pattern consists of amorphous, fl uorescent blotches, disrupted by non-fl uorescent tents. The second pattern consists of discontinuous, evenly distanced, spiral lines. The spiral lines are a series of alterations of fl uorescent dots-dashes and non-fl uorescent dashes. The second pattern, when overlapping the non-fl uorescent tent level, displays only the fl uorescent dots-dashes of the continuous spiral lines (Fig. 19).

Remarks
This species comprises only one specimen, partly broken (Table 10). Its biconical morphology is unique in the Greek collection. It could be compared with those studied by Davoli (1972) and named as ?Conus clavatulus d'Orbigny, 1852 (MOD n°5569 bis and MOD n°5610). These specimens have not been studied under UV light; therefore, we place them within Conus (Lautoconus) sp. 5. The specimen (AMPG(IV) 3861) is similar t o several species of Conus described from the Paratethys, discussed in Harzhauser & Landau (2016) as Leporiconus Iredale, 1930. This specie s differs from the Conus (Leporiconus) in lacking tuberc les on the early spire whorls. Also, the shell is smooth, except for the

Stratigraphic range
?Tortonian of Italy from Montegibbio (Davoli 1972) and Greece (Messara Basin, Crete).   Table 18). Conus (Lautoconus) sp. 1 is a species found only in Crete. Finally, this species assemblage suggests a much stronger relationship with the Proto-Mediterranean faunas than with the Paratethyan faunas.
The PCA graph (Fig. 20) shows that the species are morphologically similar and point out the necessity of the colour patterns as an extra characteristic for their identifi cation (Psarras et al. 2021

Descriptive comments
The nodules on the spire whorls occur on most extant species, except on Conus (Stephanoconus) genuanus Linnaeus, 1 758 and can either fade out or persist in later spire whorls. Some species also have spiral ribs on the body whorl. The protoconch is multispiral. The colour pattern consists of spiral bands (two to three), axial blotches, spiral rows of small to large dots and dashes, in colours of shades brown, cream and white. All Conus (Stephanoconus) feed on Polychaeta have a similar radular morphology (Tucker & Tenorio 2009).

Remarks
According to Puillandre et al. (2015), the subgenus occurs in all tropical seas.

Shell description
Usually small to rarely medium-sized shells (SL max.: 52.5 mm) (Fig. 22). Spire elevated, conical. Outline of spire whorls variable in shape, with height, ranging from medium to high (Fig. 22). Early spire whorls elevated, convex, striate, creating straight to highly conical outline, tuberculated, with faint spiral cords on early sutural ramps. Later spire whorls smooth, elongate, convex, not always in symmetry to surrounding spire whorls, with late spire whorls usually infl ated and more robust. Suture incised, slightly undulated. Subsutural fl exure shallow, moderately curved, moderately asymmetrical (Fig. 40K). Shoulder smooth to weakly angular, with maximum diameter just below shoulder. Last whorl elongated, curved. Aperture narrow, widening slightly towards fasciole. Siphonal canal short, straight. Siphonal fasciole indistinct. Faint spiral grooves visible on one fi fth anterior part of body.

Description of colour pattern
The colour p attern consists of two levels of colouration. The fi rst one is a pattern of irregular blotches (Fig. 23) of variable size, sometimes with zig-zag boundaries (Fig. 22B) or non-continuous spiral bands on the middle and anterior parts of the shell (Fig. 22C, F, H). The pattern might continue on spire whorls as irregular, thin, fl ammulated blotches (Fig. 24). Th e second level consists of evenly arranged, spiral rows of wavy dashes and dots (Figs 23-24). The spiral rows start at the shoulder of the shell, continuing spirally towards the anterior part. The spiral rows exists in between the spiral grooves. On some specimens of exceptional preservation, the spiral lines of dashes and dots are better displayed as continuous fl uorescent wavy lines, with interruptions of grey coloured oval dots (Fig. 24). These dots are visible even under natural light (Figs 21A, C, 23). When the lines cover the fi rst pattern of irregular blotches, the second pattern is seen as bright-fl uorescent (Fig. 24), while the grey coloured oval dots are often presumed to be part of the base colour, which is not the case. Spiral rows are often not visible in shells of no exceptional preservation.

Remarks
The Cretan specimens are very similar to Conus taurinensis Bellardi & Michelotti, 1841. To our knowledge, the type specimens of Conus taurinensis Bellardi & Michelotti, 1841, discussed by Sacco (1893b, Hall (1966) andFerrero Mortara (1984), are stored in the Museo Regionale di Scienze Naturali, Turin, Italy (Annalaura Pistarino pers. comm.) (syntype MRSN BS.038.05.163), but no colour patterns are preserved. Hall (1966) stated that the type specimens of Conus taurinensis possess spiral cords on their last whorl. However, these specimens are very worn. Due to the bad preservation and the lack of colour patterns of the type material of C. taurinensis, we refrain from attributing with certainty the Cretan specimens to this species until more topotypes with colour patterns are observed.  Landau et al. (2013: pl. 12 fi gs 6-7) also fi gured one Turkish specimen as Varionoconus taurinensis (Bellardi & Michelotti, 1841), this time with a visible colour pattern which is identical to that of the Cretan specimens. We consider the specimens of both Erünal-Erentöz (1958) and Landau et al. (2013) as conspecifi c to Conus cf. taurinensis.

Shell description
Small shells (SL max.: 24.17 mm). Early spire whorls elevated, convex to conical, tuberculated. Later spire whorls smooth, straight to convex, with channelled suture. Spire whorl outline conical to slightly coeloconoid in early spire whorls, conical to convex in later spire whorls. Subsutural fl exure shallow, moderately curved, moderately asymmetrical (Fig. 40L). Shoulder smooth, rounded to slightly angulated, with maximum diameter right below shoulder. Aperture of narrow to medium width, widening towards shell's anterior part. Last whorl straight to slightly widened below shoulder. Fasciole short, slightly twisted. No spiral grooves on last whorl.

Description of colour pattern
On the spire whorls, the colour pattern consists of irregular, bright-fl uorescent fl ammulae. On the last whorls, the fi rst colour pattern level consists of three, widely spaced, fl uorescent spiral bands. The bands exist near the shoulder, at the middle and anterior parts of the last whorl. The second level consists of very closely arranged, spiral lines of dashes, which can vary from very long to dot-like dashes (Fig. 27). In between the dashes, non-fl uorescent dashes exist, usually creating axially to diagonally spaced, nonfl uorescent areas, in the form of blotches. The intensity of the fl uorescent colours is not constant in the levels of patterns (Fig. 28). The fi rst level is illuminated with a fade-fl uorescent colour. The colour intensity of the second level is axially directed, from fade-fl uorescent to bright-fl uorescent in colour, creating axially arranged, bright-fl uorescent areas of dashes, suppressed by non-fl uorescent blotches. The non-fl uorescent dashes, between the fl uorescent ones, are visible when the second pattern overlaps the fi rst.

Remarks
This species is identifi able from the rest of the studied specimens in the small size of the shells and the intricate colour pattern variation of dashes and bands. Conus (Stephanoconus) c f. taurinensis is distinguishable from this species by the early highly conical spire whorls, the smoother spire whorl outline, with less infl ated sutural ramps and a different colour pattern. The shell shape of Conus (Stephanoconus) moissettei sp. nov. resembles a specimen of Conus (sensu lato) vindobonensis Hoernes & Auinger, 1879 (Harzhauser & Landau 2016: fi g. 35e and discussion), but the Austrian species differs by a more elongated shell shape, a deeper subsutural fl exure and spiral grooves on the anterior part of last whorl. Its colour pattern is not known.

Concluding remarks about Conus (Stephanoconus) Mörch, 1852
Here, this subgenus is id entifi ed in the Miocene of Europe for the fi rst time. Three species are assigned to this subgenus: Conus (Stephanoconus) asterousiaensis (Psarras & Koskeridou & Merle, 2021)

Remarks
Tucker & Tenorio (2009) fi rst described Plagioconus at genus level and were followed by Harzhauser & Landau (2016), who, in turn, proposed an emended description, according to their Paratethyan species. On the other hand, Puillandre et al. (2014,2015) placed many genera of Conidae as subgenera under the genus Conus. Morphologically, this group of species can be included in the genus Conus, thus, we refer here to Plagioconus at the subgeneric level. Michelotti, 1847 Figs 29, 36, 40O;

Shell description
Medium-large-sized shells (SL max.: 82.8 mm). Spire whorls of medium height, conical to convex in outline in early spire whorls, coeloconoid in late spire whorls. Early spire whorls conical, convex, slightly elevated. Middle spire whorls convex, with straight to convex sutural ramp. Last two to three whorls widened, with straight or infl ated, deep suture. Absence of ornamentation along the sutural ramp. Subsutural fl exure deep to very deep, rarely moderately deep, usually moderately to rarely strongly curved, strongly asymmetrical (Fig. 40O). Shoulder smooth, with weak angulation near external part of sutural ramp, more angulated on last whorl. Aperture straight, relatively narrow. Fasciole carved by growth lines. Last whorl straight conical.

Description of colour pattern
The colour pattern faintly exists in some shells in the form of axial fl ammulae in the spire whorls, parallel to the subsutural fl exure. On the last whorl, some shells present continuous axially arranged stripes united with the fl ammulae of the spire whorls. Due to the bad preservation of the specimens, some partially preserve their stripes in the form of axial blotches (Fig. 29H2).

Remarks
Michelotti (1847) identifi ed two morphotypes of Conus for which he gave two names, elatus and puschi respectively. The two species were differentiated mainly by the presence or absence of a widened last whorl and the angulation at shoulder. This difference was later illustrated in Sacco (1893b) and by Hall (1966). Davoli (1972) fi gured some extreme morphs of Conus elatus, but he failed to identify Conus puschi, as noted by Harzhauser & Landau (2016). Additionally, morphotypes of the Paratethyan Conus puschi possess shallower subsutural fl exures than Conus elatus (Harzhauser & Landau 2016). Taking all this into account, we tried to identify our specimens based on morphology, as the colour patterns are poorly preserved or non-existent. The material mostly comes from one locality (Achladhia) and was previously assigned to Conus puschi (Symeonidis 1965;Symeonidis & Kostantinidis 1968). We identifi ed several large sized specimens with a widened shoulder and angulations at the shoulder (Fig. 29F-G), which can be considered as 'elatus' morphs. We also found shells with smooth spire whorls and almost no angulation at the shoulder (Fig. 29A), which can be considered as 'puschi' morphs. Due to the large number of specimens, we were able to see their morphological variability (Table 13). We identifi ed some shells with intermediate morphologies of smooth dome-shaped spire whorls (Fig. 29B-C) and widened shoulder (Fig. 29D) and others with an angulated but normally arranged shoulder (Fig. 29H). Furthermore, the subsutural fl exure was variable in individual shells, and some had shallower SSF than others. With this evidence, we consider the two morphs being the extremes of a single species.
The Cretan specimens could be compared to Conus marii Sacco, 1893 but they differ from the fi gured specimens of Conus marii illustrated by Hall (1966) and Harzhauser & Landau (2016) in the elevated and less infl ated spire whorls.
Some of the studied specimens ( Fig. 29A-D)

Shell description
Medium-large-sized shells (SL max.: 76.1 mm). Spire whorls medium to high, with conical outline. Protoconch multispiral. Early spire whorls conical, slightly elevated and beaded, with angulated shoulder. Late spire whorls conical, fl at to striated, with straight to convex sutural ramp. Suture incised, of variable depth. Subsutural fl exure moderately deep (Fig. 40P) to deep (Fig. 40Q), moderatel y curved, strongly asymmetrical. Sutural ramp straight to slightly infl ated and convex. Shoulder smooth to subangulated. Spiral angulation at shoulder, as separation of last whorl and sutural ramp. Aperture straight, narrow. Last whorl elongated, conical. Fasciole smooth, elongated, slightly twisted. Faint spiral cords at anterior part of last whorl.

Description of colour pattern
The fi rst pattern level consists of an alteration of fl uorescent spiral bands, on a non-fl uorescent base colour, with one thin fl uorescent band at the position of the maximum diameter, another wide fl uorescent band near the shoulder of the shell, and, in some shells, a third smaller band present on half of the anterior part of the last whorl (Fig. 30B). The second level consists of rows of dots and dashes, which are axially or spirally arranged on the surface of the shell, on top of the fi rst fi eld. The colour pattern of the spire whorls consists of fl uorescent fl ammulae of irregular width, usually continuing on the last whorl, as axial dots or dashes (Fig. 30G). The pattern is not consistent in all shells, as some shells under the UV light display either faded patterns (Fig. 30C), one pattern on the last whorl (Fig. 30B), or none at all (Fig. 30D).

Remarks
The Cretan specimens are mostly broken, but the remaining shell surfaces are in pristine condition, the beads on the spire whorls are visible and, most importantly, the colour patterns can be distinguished under UV light (Fig. 31). The specimens of Conus (Plagioconus) sp. display variable shoulder angulation height of the spire ( Fig. 30; Table 14). The colour pattern is consistent, with fl uorescent bands and occasional spiral or axial dots and dashes. Some of the studied specimens have previously been identifi ed by Symeonidis & Kostantinidis (1968)      As we had no access to their colour patterns, we cannot check the synonymy of Hall (1966). A comparison with the syntype (MNHN.F.A13061) of Conus (Plagioconus) aquensis (Fig. 32) from the early Miocene of the Aquitaine Basin shows that Hall's specimens possess more angulated shoulders, high conical spires and relatively narrower diameters (RD). Two specimens named as Conus puschi from Davoli (1972: pl. 8 fi gs 17a-b, 19) have an elongated spire similar to that of the studied specimens ( Fig. 30B1-C2), but the spire whorls are not distinctly elevated and there is a slight angulation at the shoulder. We also wait for their colour pattern descriptions in order to compare to the Cretan specimens.

Shell description
Moderately large (SL max.: 67.4 mm), elongated shell. Early spire whorls beaded, elongated, with angulated shoulder. Beads absent after the 7 th spire whorl, angulated shoulder smoother after the 5 th spire whorl (Fig. 34). Delicate spiral cords on sutural ramp (Fig. 32). In some specimens, up to three spiral grooves can be seen, just under shoulder (Fig. 34). Maximum diameter just below shoulder. Suture incised, subsutural fl exure shallow, moderately curved, strongly asymmetrical (Fig. 40). Last whorl straight to slightly infl ated. Surface of subadult shells decorated with spiral grooves on all of shell's length, deeper grooves at anterior half of shell ( Fig. 34). Spiral nodules present along last whorl in between spiral grooves (Fig. 32B1), fading out on adult specimens (Fig. 32A1). Also, on adult specimens, spiral grooves present only along half to two thirds of anterior part of last whorl. Aperture narrow, straight, widened slightly on fasciole. Siphonal canal medium. Fasciole indistinct, slightly twisted.

Description of colour pattern
The colour pattern consists of fl ammulae and blotches on the spire whorls, sometimes aligned with the subsutural fl exures. Flammulae continue until just below the shoulder. The colour pattern on the last whorl consists of fl uorescent, densely spaced dots and dashes, organized axially and spirally. Fluores cent axial blotches also exist, uniting several dashes axially (Fig. 35). These fl uorescent blotches are mainly present on the upper, middle and bottom parts of the last whorl, organised as faint spiral bands.

Remarks
The Cretan specimens are very similar to the syntypes of Conus (Plagioconus) aquensis d'Orbigny, 1852 originally described from the early Miocene of Aquitaine Basin, as they share similarly beaded early spire whorls, spiral grooves along the two thirds of the anterior part of the last whorl. The subsutural fl exures are shallow and the colour pattern is identical. The species is easily recognised by the prominent beaded early to middle spire whorls (see Fig. 32B7), the grooved last whorl and the moderately elevated   (Table 15). The colour pattern of spiral dashes is also easily recognisable under UV light. Conus ( Plagioconus) aquensis d'Orbigny, 1852 has been mistakenly named as Conus antidiluvianus by Grateloup (1847) from which it mainly differs in the convex spire whorls and the colour pattern (Janssen et al. 2014). d'Orbig ny (1852) noticed the error and named this species Conus aquensis. Peyrot (1930) was the fi rst who gave a detailed description of this species and fi gured some examples. The typ e specimen of Conus burdigalensis Mayer, 1858(from Saucats near Bordeaux, Burdigalian, Mayer-Eymar coll. N 2741, fi gured in Hall (1966: pl. 27 fi g. 11) is similar in morphology to Conus (Plagioconus) aquensis, but we have not studied the specimen's colour patterns. Landau et al. (2013: Fig. 34. C onus (Plagioconus) aquensis d'Orbigny, 1852 from the Tortonian of Filippi, Crete (Greece) (AMPG(IV) 3915), specimen in close-up. A1. Notice the beaded spire whorls, the three spiral cords on shoulder and the spiral grooves on the rest of the shell. A2. Notice the spiral grooves on last whorl. A3. Notice the delicate spiral cords on sutural ramp, as well as shallow subsutural fl exure. Scale bars: A1, A3 = 2 mm; A2 = 1 mm.
pl. 81 fi g. 9) identifi ed one of their fossils as Plagioconus puschi (Michelotti, 1847) (RGM 777 891 (ex JvdV collection), Pınarlar Yaylası, Akpınar, Karaman Basin, Turkey). The colour pattern resembles the pattern described herein. The morphological characteristic of multiple spiral grooves on the anterior part of the shell are evident in their fi gure, they match those of Conus (Plagioconus) aquensis. For these reasons, we consider the Turkish specimens as Conus (Plagioconus) aquensis. Furthermore, Landau et al. (2013) compared a specimen from the Langhian of Romania attributed by Caze (2010) to Conus (Chelyconus) sp. (Caze 2010: fi g. 5m1-2), which has a similar colour pattern. We studied this specimen (MNHN.F.A31836) and confi rmed that the pattern is similar to Conus (Plagioconus) aquensis. The morphology of the spire is very similar to that of the type specimen, with beads on early spire whorls and a shallow subsutural fl exure. The pattern is also similar to Conus (Plagioconus) aquensis, but the faint spiral and axial bands, underlying the spiral dashes, cover a larger portion of the shell. This pattern is considered here as a variation; therefore, the specimen (MNHN.F.A31836) is considered here as Conus (Plagioconus) aquensis. Harzhauser & Landau (2016) named a new species Conus (Plagioconus) bellissimus, but they showed the pattern of only one specimen out of four fi gured therein (fi g. 31c-f). The shells fi gured (Harzhauser & Landau 2016) have a similar morphology to each other, except for  a specimen from the L anghian of Letkés in Hungary (private collection Anton Breitenberger), which is provided with its colour pattern, that has a more infl ated sutural ramp and a shorter spire than the rest of the specimens shown by the authors (Harzhauser & Landau 2016). This specimen illustrated under UV light bears an outline and a colour pattern similar to that of specimen AMPG(IV) 3909, Achladhia, Crete. Additionally, the colour pattern of this Hungarian specimen is identical to that of the Turkish and Romanian ones illustrated by Landau et al. (2013).

Diagnosis
A small-sized, torpedo-shaped species with delicate pattern of spiral rows of dots and blotches.

Etymology
A species named after Davoli, who fi rst fi gured that species.

Description of colour pattern
The pattern consists of small fl uorescent blotches, and evenly arranged fading spiral lines bearing dots at almost equal distances from each other (Fig. 38).

Remarks
Conus davolii sp. nov. is a characteristic torpedo-shaped species (Fig. 37), which has a very delicate colour pattern. Its morphology is unique in the Greek collection (Table 16). Two Italian specimens from the Tortonian of Montegibbio have been misidentifi ed as Conus clavatulus d'Orbigny 1852 by Davoli (1972) (see remarks for Conus (Lautoconus) sp. 3). On the other hand, their rare torpedo-shaped morphology is similar to that of the Cretan specimen. Despite the fact that we did not study the patterns of the two specimens, the Italian specimens and the Greek specimen are morphologically conspecifi c. In view of this deduction and because of lack of more material from the type locality, we designate the two specimens fi gured by Davoli (1972: pl. 4 fi gs 5, 17) as the paratypes.
This species could be misinterpreted as a juvenile Conus (Lautoconus) ictini sp. nov. The morphology of this species is different from Conus (Lautoconus) ictini sp. nov., as the spire whorls are not straight, because there are no spiral grooves on the anterior part of the last whorl. The colour pattern is also different from Conus (Lautoconus) ictini sp. nov., as there is no brick-like pattern, but a dotted pattern on the spiral lines. The species could be compared to Conus praelongus Hoernes & Auinger, 1879, but, morphologically, the spire of Conus davolii sp. nov. is not as elevated as in Conus praelongus (see Harzhauser & Landau 2016: fi g. 34b-d). The colour pattern is different as well, with Conus praelongus bearing a dashed pattern.

Conus olivaeformis
Hoernes & Auinger, 1879 is another species with a similar morphology. It differs in the shallow subsutural fl exure, the subangulated shoulder and the inferred (Harzhauser & Landau 2016) colour pattern of densely spaced spiral dashes. The specimen of Kovács & Balázs (2015) is probably Conus olivaeformis, but without UV light results, we cannot be certain. Conus davolii sp. nov. resembles the extant Conus (Hermes) nussatella Linnaeus, 1758, in the torpedoshaped whorl and the colour pattern of spiral rows of continuous dashes and spots on them, with alternating blotches. The ridges on the last whorl are a feature not present on the studied specimen; therefore, we do not assign the studied species to Conus (Hermes) Montfort, 1810 and do not include this species to any subgenus until more material will have been studied.

Description of colour pattern
The colour pattern on the last whorl consists of evenly spaced spiral rows of continuous lines. Between the spirals, very thin, wavy, mostly discontinuous spiral lines exist. The spiral rows, of continuous lines on spiral grooves, are just above every spiral groove. On the spire whorls, the spiral lines start near the shoulder of the shell, while the spiral line near the suture is partly covered by the growing shell. The lines are not always of the same width and vary from clear lines with specifi c boundaries to wavy, blurry lines.

Remarks
Harzhauser & Landau (2016) described the morphological variation of Conus fuscocingulatus and briefl y characterised its colour patterns. We consider that our material belongs to this species and expand the colour pattern description. Our specimens show a diversity at the angulation (Fig. 39A, E) and width (Fig. 39D, F) of the shoulder, as well as at the height of the spire (Fig. 39B, G) (Table 17). Nevertheless, other characteristics such as the dense spiral cords and grooves on the anterior part of the last whorl, the shallow subsutural fl exure (Fig. 40N), and the very clear colour pattern are indicative of the conspecifi city of the specimens. Usually, the spire whorls of the specimens are eroded and only few beads and indistinct swellings are visible. From those characteristics, w e could place this species in Conus (Stephanoconus) Mörch, 1852, but the colour pattern variation is clearly different from the patterns seen in extant Conus (Stephanoconus). We also disagree with the placement of this species in Conus (Phasmoconus), as this subgenus engulfs many species with radically different morphological and colour pattern variations (e.g., see Monnier et al. 2018). An extant species from Papua-New Guinea and the Solomon Islands, attributed to Conus (Phasmoconus) aff. mucronatus Reeve, 1843 by Monnier et al. (2018) displays a rather similar colour pattern, but differs from Conus fuscocingulatus Hörnes, 1851 by fl ammulae on its spire whorls, many discontinuous spiral rows and an angulate shoulder.

Discussion
This work has focused on the su bgenera Lautoconus Monterosato, 1923, Stephanoconus Mörch, 1852, and Plagioconus Tucker & Tenorio, 2009 as well as on two species of Conus not included in a subgenus. We found 17 species from the Tortonian of Crete, Greece (Table 18) (Table 18). The Cretan assemblage shares four species with the Serravallian fauna of Turkey. With the middle Miocene of Central Proto-Mediterranean and the Atlantic, it shares at least three species and with the Paratethyan region it shares at least fi ve species. At least three species seem to persist in the Pliocene of the Mediterranean (Table 18). Psarras et al. (2021) studied the Conus (Kalloconus) and Conilithes and previously identifi ed 11 species in the Tortonian of Crete, one of which was endemic to Crete. The presence of at least two more species  as it either displays endemic species from Crete, or it indicates a taxonomic gap in the inventory of the Proto-Mediterranean Conidae. In essence, the description of residual colour patterns of the Conidae is crucial for a more precise assessment of the conid paleobiodiversity.