The earliest cowries: the origin of cypraeoid gastropods

ALEXANDER NÜTZEL, SIMON SCHNEIDER, SOFIA BAKAYEVA, and ANDRZEJ KAIM

Nützel, A, Schneider, S., Bakayeva, S., and Kaim, A. 2025. The earliest cowries: the origin of cypraeoid gastropods. Acta Palaeontologica Polonica 70 (2): 213–223.

Cowries, the family Cypraeidae, form a diverse and conspicuous group of gastropods living in tropical to subtropical seas. Their shell is convolute (last whorl covers all previous ones) with a narrow, slit-like siphonate aperture bearing denticles (“teeth”). When extended, a large part of their shell surface is covered by a soft fleshy mantle. The earliest cowries were reported from the Upper Jurassic of Sicily: Cypraea tithonica and Cypraea gemmellaroi. Subsequently, these species had been assigned to various cypraeid genera. Examination of the type material of Cypraea tithonica reveals that this species represents a new genus: Coffeacypraea Nützel & Schneider. Cypraea gemmellaroi also belongs to this new genus and is potentially synonymous with Coffeacypraea tithonica. The Upper Jurassic caenogastropod genera Colombellina and Zittelia (family Colombellinidae) also have narrowly elongated siphonate apertures and are closely related to Cypraeidae but their shells are not convolute. The origination of Cypraeidae and Colombellinidae contributed considerably to the Mesozoic–Cenozoic caenogastropod radiation.

Key words: Gastropoda, Cypraeoidea, Jurassic, taxonomy, systematics, evolutionary relationships, Sicily, Italy.

Alexander Nützel [nuetzel@snsb.de; ORCID: https://orcid.org/0000-0002-8852-7688 ], SNSB-Bayerische Staatssamm­lung für Paläontologie und Geologie, Richard-Wagner-Str. 10, 80333 München (Germany); Department of Earth and Environmental Sciences, Palaeontology and Geobiology, Ludwig-Maximilians-Universität München, Richard-Wagner-­Str. 10, 80333 München, Germany; GeoBio-Center, Ludwig-Maximilians-Universität München, Richard-Wagner-Str. 10, 80333 München, Germany.

Simon Schneider [simon.schneider@casp.org.uk; ORCID: https://orcid.org/0000-0001-6493-357X ]; CASP, West Building, Madingley Rise, Madingley Road, Cambridge, CB30UD, UK.

Sofia Bakayeva [sofiyabakayeva@gmail.com; ORCID: https://orcid.org/0000-0001-9292-7652 ], Institute of Paleobio­logy, Polish Academy of Sciences, ul. Twarda 51/55, 00-818 Warsaw, Poland; State Museum of Natural History, National Academy of Sciences of Ukraine, Teatralna st. 18, Lviv 79008, Ukraine.

Andrzej Kaim [kaim@twarda.pan.pl; ORCID: https://orcid.org/0000-0001-6186-5356 ], Institute of Paleobiology, Polish Academy of Sciences, ul. Twarda 51/55, 00-818 Warsaw, Poland.

Received 1 February 2025, accepted 10 March 2025, published online 22 April 2025.

Copyright © 2025 A. Nützel et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Introduction

Cowries, the family Cypraeidae Rafinesque, 1815, comprise approximately 400 living species and subspecies, and have a rich Cenozoic and sparse Cretaceous fossil record (e.g., Schilder and Schilder 1971; Lorenz 2017, 2018; MolluscaBase 2025). Two species of latest Jurassic (Tithonian) age, Cypraea tithonica and C. gemmellaroi, were first described from Sicily (Italy) by Di Stefano (1882) and later also reported from coeval strata on Capri (Italy) (Parona 1919); they represent the earliest cowries known to date. The present contribution revises the taxonomy and systematic placement of these early cypraeids, based on re-study of the type series of Cypraea tithonica Di Stefano, 1882.

Cowries are conspicuous in having a convolute shell, with the last whorl enclosing all previous whorls and forming a narrowly elongate, slit-like aperture with anterior and posterior canals. Cypraeids can cover their entire shell with a fleshy mantle, which enables them to produce a spotlessly smooth, glossy outer shell surface. While most of the modern diversity of Cypraeidae is located in the tropical Indo-Pacific, the family as a whole has a pantropical distribution between 35° N and S (Riedel 2000; Passamonti 2015; Lorenz 2017, 2018).

Juveniles of the type genus of the family, Cypraea, are herbivorous grazers, and are carnivorous or omnivorous grazers as adults, feeding on sponges and other sessile invertebrates (Tay et al. 2023 and references therein). All species of which early ontogeny and protoconchs are recorded have planktotrophic larval development. They have relatively large larval shells, generally approximately 1 mm high, but some reaching up to 1.5 mm, with a reticulate ornament (Bandel et al. 1997; Riedel 2000; Tay et al. 2023) (Fig. 1).

Cypraeidae have a taenioglossate radula and were placed in the traditional Mesogastropoda (e.g., Wenz 1938–1944). They are now assigned to the “higher” Caenogastropoda within Latrogastropoda (e.g., Bouchet et al. 2017; Ponder et al. 2020; Li et al. 2024), which are characterized by an inhalant flow, commonly associated with the development of an anterior siphonal canal. While family-level phylogeny and systematics of the Cypraeidae within Gastropoda is relatively well resolved and stable (Meyer 2003, 2004), subdivision at the genus level is still under debate. So far, cypraeid genus-level systematics is largely based on shell morphology, which is rather conservative in this family, and modern anatomical and molecular studies are lacking (e.g., Passamonti 2015). More than 150 genus-level names have been proposed for extant and fossil Cypraeidae, and approximately 80 of these are currently considered valid (Schilder and Schilder 1971; Riedel 2000; MolluscaBase 2025).

The aim of this study is to revise the available cypraeid materials from the Upper Jurassic of Italy and attempt to relate them to other caenogastropods which have narrowly elongated siphonate apertures.

Institutional abbreviations.—MGUP, Museo Geologico G.G. Gemmellaro, Università di Palermo, Sicily, Italy; MSTC, Museo di Paleontologia, Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Università di Catania, Sicily, Italy; SNSB-BSPG, Bayerische Staatssammlung für Paläontologie und Geologie, München, Germany.

Nomenclatural acts.—This published work and the nomenclatural acts it contains have been registered in Zoobank: urn:lsid:zoobank.org:pub:3FC4C848-4D6B-4554-AB84-1EB2125A6D01.

Historical background

As highlighted above, the earliest Cypraeidae have been described from the uppermost Jurassic of Sicily (Di Stefano 1882; Tracey et al. 1993; Groves and Landau 2021; Capasso 2024). However, the type material of Cypraea tithonica Di Stefano, 1882, and Cypraea gemmellaroi Di Stefano, 1882, has never been restudied, and Zamberlan and Checchi (2014) and Capasso (2024) considered these specimens as lost. Thus, it came as a surprise that during a visit to the Museo di Geologia G.G. Gemmellaro in Palermo, two of the authors (AN and SS) re-discovered the type series of Cypraea titho­nica, which forms the base for the present contribution. The original material of Cypraea gemmellaroi was not found.

Besides their type locality, Termini Imerese near Paler­mo, both species were reported to occur in Tithonian strata on the island of Capri south of Naples, Italy by Parona (1919); as far as we are aware, his specimens were never illustra­ted. However, Zamberlan and Checchi (2014) figured a speci­men from the Tithonian of Capri, which is part of the Viglino Collection (MSTC no. 186vi0950) curated at the Museo di Paleontologia, Dipartimento di Scienze Bio­logiche, Geologiche e Ambientali, Università di Catania, and was assigned to Cypraea gemmellaroi; they further stated that the same collection also contained specimens of Cypraea titho­nica (MSTC no. 186vi0949). Recently, Capasso (2024) figu­red both specimens from Capri.

Cypraea tithonica was assigned to Cyproglobina De Gregorio, 1880, by Schilder (1927), a genus now placed in the Ovulidae (Fehse 2013). Later, Schilder and Schilder (1971) transferred the species to Palaeocypraea Schilder, 1928, where it was retained by Groves (1990, 1992, 1994), Tracey et al. (1993) and most recently Capasso (2024). Based on re-study of the type material of C. tithonica, this assignment is erroneous, as will be outlined below.

The second, larger species introduced by Di Stefano (1882), Cypraea gemmellaroi, was assigned by Schilder (1927) to his new subgenus Bernaya (Protocypraea), but later transferred to Bernaya Jousseaume, 1884 sensu stricto (Schilder and Schilder 1971). Again, Groves (1990, 1992, 1994), Zamberlan and Checchi (2014) and Capasso (2024) retained this combination. This assignment is equally untenable, and Cypraea gemmellaroi is congeneric with Cyp­raeatitho­nica.

Material and methods

The material studied herein is housed at the Museo Geo­logico G.G. Gemmellaro in Palermo, Sicily, Italy. Only the type series of Cypraea tithonica Di Stefano, 1882, consisting of seven syntypes, which are archived under accession numbers MGUP-20-368a–g, was found during a visit in March 2017. The type series of Cypraea gemmellaroi Di Stefano, 1882, could not be traced.

The specimens come from an isolated limestone outcrop on the castle mountain of Termini Imerese (Palermo region, Sicily; coordinates: 37.9886, 13.6979, WGS1984). The castle mountain succession is part of a series of originally African structural domains (Sicilian-Maghrebide units; Fig. 2) that were obducted onto the European part of Sicily during Neogene times (Catalano et al. 2010, 2011). The strata that yielded the cypraeids are assigned to the informal Ellipsactinia breccias member of the Crisanti Formation (Catalano et al. 2010; Basilone 2018), which comprises thick-bedded limestone breccias and conglomerates. The components of these rocks are characterised by shallow-­water carbonate platform facies containing abundant corals and molluscs; the latter were monographed by Gemmellaro (1869a–d, 1871a–d, 1875) and Gemmellaro and Di Blasi (1874). The rocks are classically attributed to the Tithonian, but may partly be Berriasian (earliest Cretaceous) in age, based on calpionellid biostratigraphy (Catalano et al. 2010; Basilone 2018).

All seven specimens were coated with ammonium chloride, and photographs from those views deemed taxonomically informative were taken. The higher classification of the Gastropoda is adopted from Bouchet et al. (2017), Ponder et al. (2020) and MolluscaBase (2025). The descriptive terminology follows Groves and Squires (2023).

In 2024, SB and AK analysed the type material of the colombellid Zittelia cipraeaeformis Gemmellaro, 1869d from the Gemmellaro collection stored at the same museum, which comprises nine syntypes (SB and AK unpublished data). All of them are stored under the inventory number MGUP-20-361.

The Recent specimens depicted in Fig. 1 are stored at the Bayerische Staassammlung für Paläontologie, München (SNSB-BSPG).

Systematic palaeontology

Class Gastropoda Cuvier, 1795

Infraclass Caenogastropoda Cox, 1960

Cohort Sorbeoconcha Ponder & Lindberg, 1997

Megaorder Hypsogastropoda Ponder & Lindberg, 1997

Superorder Latrogastropoda Riedel, 2000

Order Cypraeida Rafinesque, 1815

Superfamily Cypraeoidea Rafinesque, 1815

Family Cypraeidae Rafinesque, 1815

Genus Coffeacypraea Nützel & Schneider nov.

Zoobank LSID: urn:lsid:zoobank.org:act:C0956AF7-1075-4294-8F6F- 14EC45D48245.

Type species: Cypraea tithonica Di Stefano, 1882, by original designation herein.

Species included: Coffeacypraea tithonica (Di Stefano, 1882) and Coffeacypraea gemmellaroi (Di Stefano, 1882).

Etymology: A combination of the Latin genus names Coffea, which includes the coffee plant, and Cypraea, which is the type genus of the Cypraeidae, referring to the rather simple shell shape of the new genus, which resembles an oversized coffee bean.

Diagnosis.—Relatively small (up to 25 mm long), smooth-shelled Cypraeidae; short-ovate to almost circular in outline in dorsal view; basal side of shell widely flattened; dorsal side more or less evenly arched and inflated. Aperture slit-like, narrow, in almost central position, straight, without widened anterior and posterior notches; columellar and labral denticles evenly sized and spaced (barely visible due to limited preservation). Anterior (siphonal) and posterior canals indistinct or very short.

Stratigraphic and geographic range.—Tithonian, Upper Jurassic of Italy and potentially Czechia.

Coffeacypraea tithonica (Di Stefano, 1882)

Figs. 3A–D, 4A–G, 5.

1882 Cypraea tithonica, Di Stef.; Di Stefano 1882: 76, pl. 4: 4, 5.

?1911 Cypraea tithonica di Stef.; Blaschke 1911: 162.

?1919 Cypr. tithonica Di Stef.; Parona 1919: 475.

1927 [Cyproglobina] tithonica di Stef.; Schilder 1927: 71.

1971 Palaeocypraea (Palaeocypraea) tithonica; Schilder and Schilder 1971: 24.

1990 Palaeocypraea (P.) tithonica (Stefano, 1882); Groves 1990: 273.

1992 Palaeocypraea (Palaeocypraea) tithonica (Stefano, 1882); Groves 1990: 101.

1993 Palaeocypraea tithonica; Tracey et al. 1993: 148.

1994 Palaeocypraea (Palaeocypraea) tithonica (Steffano 1882) [sic!]; Groves 1994: 26, figs. 5, 6 [reproduced from Di Stefano 1882].

2014 Bernaya gemmellaroi (Di Stefano, 1882); Zamberlan and Checchi 2014: 21, fig. 8 [not fig. 7].

2021 Palaeocypraea tithonica (Stefano, 1882); Groves and Landau 2021: 8.

2024 Palaeocypraea tithonica (Di Stefano, 1882); Capasso 2024: 165, figs. 1, 2.

2024 Bernaya gemmellaroi (Di Stefano, 1882); Capasso 2024: 166, fig. 4 [not fig. 3].

Emended diagnosis.—Small (up to 13 mm long), smooth-shelled Cypraeidae; short-ovate to almost circular in outline in dorsal view; basal side of shell widely flattened; dorsal side more or less evenly arched and inflated. Aperture slit-like, in almost central position, straight, without widened anterior and posterior notches; columellar and labral denticles evenly sized and spaced (barely visible due to limited preservation). Anterior (siphonal) and posterior canals very short.

Material.—Seven syntypes, curated under catalogue numbers MGUP-20-368a–g; relatively well preserved shells. The specimen in Fig. 4D (Di Stefano 1882: pl. 4: 4a, b; MGUP-20-368a) is designated as the lectotype herein, while the remaining six specimens become paralectotypes (MGUP-20-368b–g). All specimens come from the castle mountain of Termini Imerese, Palermo Region, Sicily; Ellipsactinia breccias member, Crisanti Formation, Tithonian, Upper Jurassic. Two additional specimens not examined by the authors come from Tithonian strata on the island of Capri (MSTC no. 186vi0949 and MSTC no. 186vi0950); see Zam­berlan and Checchi (2014) and Capasso (2024).

Description.—Shell up to 13 mm long, 12.5 mm wide and 8 mm high; convolute, cypraeid in shape, with last whorl overgrowing all previous whorls; very short ovate to almost circular in outline in dorsal view; moderately and evenly inflated in lateral view, with basal (apertural) side flattened. Outer shell surface smooth. Aperture narrow, slit-like, in almost central position, straight, without widened anterior and posterior notches. Columellar and labral sides of aperture with numerous teeth, in most specimens barely visible due to limited preservation.

Remarks.—The studied specimens of Coffeacypraea titho­nica generally are similar to modern cypraeids with regard to shell morphology, but differ sufficiently from younger taxa to be placed in their own genus. The specimens illustrated in Fig. 4A and B show that the last whorl overgrows the low-spired previous whorls and produces the typical cypraeid convolute shape. However, as noted by Capasso (2024), the shells are much shorter and more circular than in any other genera, with length/width ratios of 1.04 to 1.18 for C. tithonica and approximately 1.12 to 1.15 for C. gemmellaroi (measured from Fig. 3E, F). Coffeacypraea Nützel & Schneider gen. nov. also differs from the other genera of the Cypraeidae by having a straight, rather narrow aperture in central position, which lacks the widened anterior and posterior notches seen in their Cretaceous and younger descendants. Moreover, the anterior and posterior canals are barely developed. Although the preservation of the specimens is not ideal, this is not a result of erosion, but a genuine feature. By far most other Cypraeidae have more or less curved apertures, which are opisthocyrt inclined and the siphonal (anterior) and posterior canals are usually much more pronounced.

Two specimens from the island of Capri, figured by Zam­berlan and Checchi (2014) and Capasso (2024) under the names Bernaya gemmellaroi (Di Stefano, 1882) and Palaeo­cypraea tithonica (Di Stefano, 1882), fall into the size range and morphological variability of the type series of C. tithonica. MSTC no. 186vi0950 is slightly tapered posteriorly and has a shallow dent in its anterior part. However, similar features occur in the paralectotypes (Fig. 4B, C, G3) and should be attributed to intraspecific variability.

Whether the single specimen from the Tithonian–Berri­asian of Štramberk (Czech Republic) reported by Blaschke (1911) should be assigned to Coffeacypraea tithonica is unconfirmed. The specimen was not illustrated and could not be traced among Blaschke’s (1911) material, which only comprises the figured specimens (personal communication Thomas Nichterl, January 2025).

In his first major revision of the Cypraeoidea, Schilder (1927) assigned Cypraea tithonica, the type species of Cof­fea­cypraea Nützel & Schneider gen. nov., to Cypro­glo­bina De Gregorio, 1880, a genus that is based on an Eocene type species from southeastern France and is presently placed in the family Ovulidae (see Fehse 2013 for subsequent type designation and taxonomy). Subsequently, Cypraea tithonica was assigned to the genus Palaeocypraea Schilder, 1928, by several authors (see synonymy above), which is based on a type species from the Danian (Paleogene) of Denmark, Cypraeacites spiratus Schlotheim, 1820. While comparable in size, Palaeocypraea spirata is significantly more elongated and has a distinctly curved aperture in a less central position. It also differs by having widened posterior and anterior canals. Owing to the curvature of the aperture, the apertural denticles vary in strength and inclination in Palaeocypraea, while they are apparently equal in size and orientation in Coffeacypraea Nützel & Schneider gen. nov. Coffeacypraea tithonica was not originally included when Schilder (1928) introduced Palaeocypraea, but was later transferred to that genus (Schilder and Schilder 1971).

Stratigraphic and geographic range.—Tithonian, Upper Jurassic of Italy and potentially Czechia.

Coffeacypraea gemmellaroi (Di Stefano, 1882)

Fig. 3E, F.

1882 Cypraea Gemmellaroi, Di-Stef.; Di Stefano 1882: 75, pl. 4: 3.

?1919 Cypraea Gemmellaroi Di Stef.; Parona 1919: 475.

1927 [Bernaya (Protocypraea)] gemmellaroi di Stef.; Schilder 1927: 88.

1928 [Protocypraea] gemmellaroi di Stef.; Schilder 1928: 23.

1971 [Bernaya (Bernaya)] gemmellaroi Stefano 1882; Schilder and Schilder 1971: 27.

1990 Bernaya (B.) gemmellaroi (Stefano, 1882); Groves 1990: 273.

1993 Bernaya gemmellaroi Stefano, 1882; Tracey et al. 1993: 148.

1994 Bernaya (Protocypraea) gemmellaroi (Steffano 1882) [sic!]; Groves 1994: 26, figs. 3, 4 [reproduced from Di Stefano 1882].

non 2014 Bernaya gemmellaroi (Di Stefano, 1882); Zamberlan and Checchi 2014: 21, fig. 8.

2021 Palaeocypraea gemmellaroi (Stefano, 1882); Groves and Landau 2021: 8.

non 2024 Bernaya gemmellaroi (Di Stefano, 1882); Capasso 2024: 166, fig. 4.

Emended diagnosis.—Relatively small (approximately 25 mm long), smooth-shelled Cypraeidae; rounded short-ovate in outline in dorsal view, with slightly tapering anterior and posterior portions; basal side of shell widely flattened, slightly subsiding towards the apertural slit; dorsal side evenly arched and inflated. Aperture slit-like, in almost central position, straight, seemingly slightly widened in central part, without widened anterior and posterior notches; apertural denticles seen on columellar lip only, presumably due to limited preservation, evenly sized and spaced. Anterior (siphonal) and posterior canals very short.

Material.—The type material of Cypraea gemmellaroi was not found during our visit to Palermo in 2017. From Di Stefano’s (1882) description, it is unclear whether there were more specimens than the figured one. However, the expression “height of the figured specimen” may be read as a hint that there were several syntypes. The type material comes from the castle mountain of Termini Imerese, Palermo Region, Sicily, Italy; Ellipsactinia breccias member, Crisanti Formation, Tithonian, Upper Jurassic.

Description.—Shell up to 25 mm long; convolute, cypraeid in shape, with last whorl overgrowing all previous whorls; rounded short-ovate in outline in dorsal view; anterior and posterior ends slightly tapering, the posterior one more so than the anterior one (note that the posterior end is facing upward in Fig. 3E, F). Shell considerably and evenly inflated in lateral view, with basal (apertural) side widely flattened, probably slightly subsiding towards the apertural slit. Outer shell surface smooth. Aperture relatively narrow, slit-like, in almost central position, slightly widened (?) in central part, straight without widened anterior and posterior notches. Apertural denticles evenly sized and spaced, clearly visible on columellar lip only, presumably due to limited preservation. Anterior (siphonal) and posterior canals very short; posterior canal more clearly marked than siphonal canal.

Remarks.—Cypraea gemmellaroi is also included in Coffea­cypraea Nützel & Schneider gen. nov., which has a more bulbous shell than Coffeacypraea tithonica and is also significantly larger, almost twice the size. According to the original illustration by Di Stefano (1882), its slit-like aperture is also straight; the anterior and posterior canals are more distinct than in Coffeacypraea tithonica but still not very pronounced. In modern cypraeids, shell size is highly variable, and in some species, the largest individuals are three times the size of the smallest ones (e.g., Poppe and Goto 1991). This means that Coffeacypraea tithonica and C. gemmellaroi may represent a single species, based on their sizes alone. However, in view of the slight differences noted above, while relying only on Di Stefano’s (1882) figures and description, both species are considered valid.

As mentioned above, Coffeacypraea gemmellaroi was assigned to Bernaya (Protocypraea) Schilder, 1927, with the original description of the subgenus (Schilder 1927; type species: Cypraea orbignyana Vredenburg, 1920; Upper Creta­ceous, southern India); Groves (1994) reverted to this assignment without discussion, but subsequently transferred the species to Bernaya Jousseaume, 1884 sensu stricto (Schilder and Schilder 1971), which is based on a type species from the Eocene of France, Cypraea media Deshayes, 1835. Species of Bernaya differ from Coffeacypraea gemmellaroi by having a curved aperture that is widened posteriorly, while that of C. gemmellaroi is straight and lacks a widened posterior part. Moreover, species of the genus Bernaya are broadly pyriform in dorso-­ventral view. In contrast, Coffeacypraea gemmellaroi is rounded short-ovate in outline. Besides the morphological differences, Eocene species of Bernaya are also significantly larger than these of Coffeacypraea Nützel & Schneider gen. nov., ranging between 37 and 62 mm in length (e.g., Pacaud 2018).

Stratigraphic and geographic range.—Tithonian, Upper Juras­sic of Italy.

Discussion

Evolutionary relationships.—The new genus Coffea­cyp­raea Nützel & Schneider in the Cypraeidae and the genera Zittelia and Colombellina in the Colombellinidae, all from the Upper Jurassic of southern and central Europe, are among the oldest caenogastropods that share an elongated, slit-like aperture, although some Paleozoic subulitoids also have narrow aper­tures. Furthermore, elongated apertures occur in several Triassic, Jurassic, and younger heterobranchs (e.g., Gründel and Nützel 2013; Hausmann and Nützel 2015). This feature has been interpreted as a convergent anti-predatory adaptation, which evolved repeatedly in the course of the Mesozoic Marine Revolution (Vermeij 1977, 1987).

With the exception of certain bellerophontoids, which are, however, isostrophically coiled, convolute gastropod shells (last whorl covering all previous ones) are unknown from the Paleozoic and the Triassic. Convolute shells with elongate aperture are also known from the Late Jurassic onward in Heterobranchia (genus Volvocylindrites Cossmann, 1895, see Gründel and Nützel 2013). So far known, Coffea­cypraea Nützel & Schneider gen. nov. represents the earliest record of this type of convolute shell in Caenogastropoda and clearly is the earliest member of the Cypraeidae and the Cypraeoidea, as was assumed previously.

The relationship between the Jurassic colombellinid Zit­telia and the Recent Cypraea was initially discussed by Sayn (1932), and later, Colombellinidae were considered the stem group of Cypraeoidea by Taylor and Morris (1988). This hypothesis, however, has never been critically assessed and Gemmellaro’s (1869d) type material had not been revised since the original description. Moreover, the characteristics of Zittelia and Colombellina have not been clearly demarcated. As a result, species have commonly been misplaced in either of these genera and taxa with entirely different characteristics were included in the family Colombellinidae (SB and AK unpublished data).

Colombellina is similar to Zittelia both with regard to its aperture morphology and its coarse nodular ornamentation (Fig. 6A, B). However, in Colombellina, the outer and inner lips are rather narrow, and the majority of the spire is not covered by them, whereas in Zittelia the lips are expanded over the spire and cover more than half of it (Fig. 6B2). This indicates that the mantle of Zittelia could envelop a significant portion of the shell. The spire, therefore, is not visible on the ventral side of the shell, as it is covered by the expansion of the lips.

According to Gemmellaro (1869d), one of the ­diagnostic characteristics of Zittelia, which distinguishes it from Colombellina, is its very narrow and slightly curved aperture, terminating both anteriorly and posteriorly in a notch or canal (Fig. 6B1). The similarity of Zittelia and Cypraea was highlighted by Gemmellaro (1869d) through the naming of one of the new species, Zittelia cypraeaeformis, which was subsequently designated as the type species of Zittelia by Cossmann (1904). Other Jurassic species of Zittelia, i.e., those described by Zittel (1873) and Brösamlen (1909) are still awaiting revision. In Cretaceous Cypraeidae, the notches seen in Zittelia have not been mentioned nor depicted (Sayn 1932; Roman and Mazeran 1920; Groves 1990, 2004; Groves et al. 2011). In the Eocene cypraeid Cypraeorbis, Darragh (2011: fig. 1) noted a similar structure, termed infracolumellar groove, located at the anterior end of the aperture (Fig. 6E1), which might be a homologous to the anterior sinuosity.

In the Late Jurassic forms of Colombellina, Zittelia, and Coffeacypraea Nützel & Schneider gen. nov., the position of the posterior canal is on the right (labral) side of the shell axis (Fig. 6A, B, C, Table 1). In Early Cretaceous cypraeids, the posterior canal has already shifted to the left (Fig. 6D), where it also occurs in Recent representatives of the Cypraeidae. It might be, therefore, hypothesized that a gradual shift of the posterior canal from the right side of the shell axis to its left side occurred between the Tithonian (Late Jurassic) and Barremian (Early Cretaceous).

Furthermore, the infracolumellar groove at the anterior end of the aperture observed in some Cenozoic and Recent Cypraeidae may have evolved from the sinuosity or notches in Zittelia and may thus represent a derived trait. However, this characteristic does not consistently appear within all cypraeids. Nevertheless, its repetitive occurrence, along with the strongly expanded lips in Zittelia, may be considered as a strong support of the relationship between these genera. Their co-occurrence in the uppermost Jurassic of Sicily, Italy, and potentially also Czechia, may indicate that the Late Jurassic reefs and associated lagoons of the Tethys were a centre of diversification of the Colombellinidae. In the same environments, decapod crustaceans underwent a remarkable diversification during Late Jurassic times (e.g., Klompmaker et al. 2013) and, as potential predators of the gastropods, may have indirectly contributed to the evolution of convolute shells with restricted apertures.

The origination of Cypraeidae with the new genus Coffeacypraea Nützel & Schneider and the diversification of Colombellinidae in the Middle to Late Jurassic contributed considerably to the Mesozoic–Cenozoic caenogastropod radiation. Both taxa represent “higher” caenogastropods (Neomesogastropoda sensu Bandel 1991 and Latrogastropoda sensu Riedel 2000). At approximately the same time, the family Purpurinidae likely gave rise to Neogastropoda (Bakayeva et al. 2024). Therefore, the Middle to Late Jurassic was a pivotal time for the evolution of Caenogastropoda, one of the most diverse animal clades.

Conclusions

Based on the restudy of the type material of Cypraea tithonica from the Tithonian of Sicily, this species is placed in the new genus Coffeacypraea Nützel & Schneider gen. nov. Coffeacypraea tithonica and Coffeacypraea gemmellaroi from the same formation represent the earliest undoubted members of Cypraeoidae. As was previously suggested, cypraeoids are closely related (possibly sister-group) to coeval Colombellinidae which also have narrow, elongate apertures but are not convolute. The origination of Cypraeidae and the diversification of Colombellinidae in the Middle to Late Jurassic contributed considerably to the Mesozoic–Cenozoic caenogastropod radiation.

Acknowledgements

This study evolved from a visit to the Museo Geologico G.G. Gem­mellaro in Palermo, Sicily, Italy, where curator Carolina d’Arpa gave us access to the collections and also valuable information on the Jurassic of the Palermo region. Lindsey T. Groves (Natural History Museum of Los Angeles County, California, USA) and Stefano Monari (Dipartimento di Geoscienze, Universitàa di Padova, Italy) are acknowledged for their careful reviews. Paolo Albano (Stazione Zoologica Anton Dohrn, Napoli, Italy) and Lindsey T. Groves kindly provided literature. Thomas Nichterl (Natural History Museum, Vienna, Austria) provided information on specimens under his care. The research of SB and AK was financially supported by the National Science Centre, Poland; grant number: 2018/31/B/ST10/03415. SB is greatly thankful for the generous support of a fellowship from the Institute of International Education’s Scholar Rescue Fund.

Editor: Eli Amson.

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Acta Palaeontol. Pol. 70 (2): 213–223, 2025

https://doi.org/10.4202/app.01245.2025