A New Genus in a Middle Jurassic Gastropod Faunula from the Vértes Mts (Hungary)

Home , Collonia, Colloniidae, Turbinidae, Turbinoidea

ANNALES HISTORICO-NATURALES MUSEI NATIONALIS HUNGARICI Volume 104 Budapest, 2012 pp. 417–430

Vertesella (Colloniidae), a new genus in a Middle Jurassic gastropod faunula from the Vértes Mts (Hungary)

J. SZABÓ

Department of Palaeontology and Geology, Hungarian Natural History Museum, H-1083 Budapest, Ludovika tér 2, Hungary. E-mail: [email protected]

– Remains of six gastropods are available from an outcrop that is one of the rare Jurassic fossil localities in the Vértes Mts. Most shells are poorly preserved; however, a single specimen indicates a new genus, Vertesella gen. n. with its type species, Vertesella vertesensis sp. n. in the faunula. Shell morphology suggests that it belongs to Colloniidae of its original diagnosis; pre-Cretaceous origin of this family seems very likely from a group of Jurassic genera, however, not all malacologists share this opinion. The occurrence of this new genus prompts us to continue thinking about the relationships and history of this insufficiently known but diversely interpreted family. With 15 figures.

– Colloniidae, Vertesella, Middle Jurassic, Vértes Mts.

INTRODUCTION

Frequency and extent of the bottom parts, suitable for gastropod ha- bitats, gradually decreased in the Hettangian to Pliensbachian (Lower Ju- rassic) interval in the whole Transdanubian Range (Hungary) to which the Vértes Mts also belong. This course dramatically accelerated in the lower- most Toarcian then with Bathonian events, which first resulted in increasing hiatuses of carbonate sedimentation then in radiolarite deposition, and ap- parently swept away (? traces of) the benthos for a long period. The Csóka- hegy gastropods belong to one of the latest Middle Jurassic benthic assemb- lages, known from the Transdanubian Range.

Annls hist.-nat. Mus. natn. hung. 104, 2012 418 J. Szabó

LOCALITY AND MATERIAL

Csóka-hegy is a hill north from the village Csókakõ, at the SW edge of the Vértes Mts next to the Móri-árok, a wide tectonic trench that separates the Bakony Mts (Fig. 1; coor- dinates of top: 47°22’14.3” N, 18°15’19.65” E). The gastropods discussed in this paper were found in 1957. Their locality cannot be identified accurately; the label of the container box indicates the SW part of the hill.

Geography of the locality. Black quadrangle: position of the area of the more detailed map within Hungary. Black asterisk indicates the outcrop(s), yielding the gastropods

Five fragmentary gastropod shells and a single, detached, calcareous operculum were collected together with a rather rich and diverse ammonite fauna by staff members from the Geological Institute of Hungary, G. VÍGH (jun.) and J. FÜLÖP. The results of the first study were published by FÜLÖP et al. (1960), then completion and corrections were given by VÍGH (1968), and FÜLÖP (1971); they identified Sinemurian (Lower Jurassic), Bajocian and Bathonian (Middle Jurassic) limestones. GALÁCZ (1995) re-studied the site, revised the former Middle Jurassic ammonite collection, and then refined the probable stratigraphical situation; he listed Late Bajocian and Late Bathonian faunas, respectively. However, he expressed some uncertainties because the “original whereabouts” of certain specimens in the studied museum material remained unreliably reconstructed. GALÁCZ (1995) recognised also that the available fossils had been preserved most probably in submarine sedimentary dykes.

Annls hist.-nat. Mus. natn. hung. 104, 2012 Vertesella, a new genus in a Middle Jurassic gastropod faunula (Colloinidae) 419

VÖRÖS &DULAI (2007) restudied the brachiopods from the same collection; they identified also Bajocian (Humphriesianum Zone) specimens. BUDAI et al. (2008) provide further information about the locality; their detailed geological map (p. 359, Plate XV: 2) shows a few dozens of usually small Jurassic spots on Csóka- hegy. They verified the presence of neptunian dykes, which penetrate beds of the Triassic Dachstein Limestone and the Fõdolomit (“Hauptdolomite”) Formations, respectively. As the latter studies demonstrate, the Csóka-hegy locality belongs to KONDA’s (1970) “discontinuous” succession type, i.e. only few sedimentary episodes, separated by signifi- cant stratigraphical gaps, are documented in condensed strata or as fissure-fillings in older rocks. This type of sedimentation was characteristic for the palaeoenvironments of submarine elevations. Gastropods remained unidentified until now; each of the listed species are represen- ted by single specimens; they are deposited in the Museum of the Geological and Geophy- sical Institute of Hungary (GGIH); the inventory numbers are: J.2012.1–6.

Subclass Archaeogastropoda THIELE, 1925 Superfamily Ataphroidea COSSMANN, 1915 Family ? COSSMANN, 1915 Ataphridae? sp. (a cross-section on rock surface), J.2012.2. Family COSSMANN, 1917 Vertesella vertesensis gen. et sp. n. (Figs 9–15), J.2012.1. Superfamily Trochoidea RAFINESQUE, 1815 Family COX, 1960 Proconulus baldensis (PARONA, 1894) (Fig. 8), J.2012.3. Subclass Caenogastropoda COX, 1959 Superfamily Loxonematoidea KOKEN, 1889 Family COSSMANN, 1909 Telleria? cf. benacensis (PARONA, 1894) (Fig. 4), J.2012.4. Superfamily Zygopleuroidea WENZ, 1938 Family WENZ, 1938 Diatrypesis angulocostatum (SZABÓ, 1983) (Figs 2–3), J.2012.5. Subclass doubtful indet. sp. (operculum) (Figs 5–7), J.2012.6.

These gastropods occurred in two lithological types; one of them is bioclastic limestone with spots of sparry calcite in the predominant micrite cement. This matrix resemb- les the characters of some varieties of the Lower Jurassic Hierlatz Limestone Formation; its lithological type occurs in small quantity also on and around the Middle Jurassic submarine heights. Its gastropod species (Diatrypesis angulocostatum, Telleria? cf. benacensis and Ataphridae? sp.) are poorly preserved. Formerly Telleria benacensis was found in the Hum- phriesianum and the Subfurcatum Zones (Bajocian) while Diatrypesis angulocostatum occurred questionably in the Humphriesianum Zone and certainly in beds of condensed Subfurcatum to Niortense Zone (CONTI &FISCHER 1984 as “Diatrypesis settepassii”; SZABÓ

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1983 as “Procerithium (Cosmocerithium)? angulocostatum”, and CONTI &SZABÓ 1987 as Diatrypesis angulocostatum). Remaining specimens are preserved with Fe–Mn-oxide coating that helped to get them in better state of preservation. Small remnants of the matrix are different from the previous limestone type; dispersed Mn-oxide content causes greyish shade of the domi- nant red colour. Proconulus baldensis (PARONA), Vertesella vertesensis gen. et sp. n. and the operculum came from this limestone. Formerly Proconulus baldensis occurred also in Bajocian strata (SZABÓ 1981, as P. rimosus,CONTI &FISCHER 1983, 1984, CONTI &SZABÓ 1987). The biostratigraphic levels of the earlier occurrences of the listed gastropod species were identified by co-occurring ammonites by GALÁCZ (1976, Somhegy, Hungary), PALLINI (1982, Case Canepine, Umbria, Italy) and STURANI (1971, Acque Fredde, Lake Garda, Italy). These data suggest that the Csóka-hegy finds have more probably come also from the Bajocian parts of the outcrops.

2 11 9 6 13

10 12 3 7

4 8 14 15 Gastropods from the Csóka-hegy (Vértes Mts, Hungary), Bajocian. 2 = Diatry- pesis angulocostatum (SZABÓ, 1983), subadult whorls (×1.5). 3 = Details of ornament in D. angulocostatum (×3). 4 = Telleria? cf. benacensis (PARONA, 1894) (×2). 5–7 = Calcareous operculum of unidentifiable species: lateral (5), inner (6) and outer view (7) (orientations to dextral shell) (×5.3). 8 = Proconulus baldensis (PARONA, 1894) (×5). 9–15 = Vertesella vertesensis gen. et sp. n., apical (9), “apertural” (10), basal (11) and abapertural (12) views (×6.7), details of peri-umbilical region (13) (×10), remnants of the original shell pattern; dark, subregularly arranged spots are preserved under the outermost, bright glaze layer; spire side (14) (×10), single row of spirally arranged spots on the base (15) (×4.2)

Annls hist.-nat. Mus. natn. hung. 104, 2012 Vertesella, a new genus in a Middle Jurassic gastropod faunula (Colloinidae) 421

SYSTEMATICS

The above sketch about the taxonomic composition of the faunula includes also not fully resolved classification problems, worthy to analyse, but they are mostly beyond the scope of the paper. This chapter aims to introduce the new genus and to discuss solely its inferred supra-generic position in the system. COSSMANN in COSSMANN &PEYROT (1917) established Colloniidae on extinct genera. The main distinctive criterion of Collonia-related shells was the lack of nacre from the shells. Besides, the definition of the family was completed also with concomitant outer shell morphological features. Most important ones were visible on the base, a more or less evident umbilicus (phaneromphalus or pseudomphalus type), and a single, circum-umbilical elevation with a terminal process on the abapical part of the columellar lip. These characters and the different morphology of the opercula provided basis to the distinction from Turbi- nidae. While the shell structure commonly changes during the fossilisation owing to re-crys- tallisation or substitution, and the opercula are normally lacking, the outer shell morphology remains more frequently accessible for study. This is why the family concept below fo- cuses on those characters of Colloniidae, which more effectively support the taxonomical practice of the palaeontology. Mainly basedon operculum characters, extant genera were also added to Colloniidae. Their shells are sometimes significantly different from those of the original family model; some of them would not be in Colloniidae by the original diagnosis. In some reinterpreta- tions of the (sub)family, the shells and especially the fossil genera are ignored. This is a rather peculiar situation in the case of a family, which was established exclusively on shells of extinct genera. In subsequent classifications, the calcified nature of the opercula and similarities in their development were mostly regarded just as indication of close relation between Collo- niidae and Turbinidae. These classifications treated Colloniidae as a subfamily within Turbinidae (WENZ 1938, KNIGHT et al. 1960, PCHELINCEV &KOROBKOV 1960, HICKMANN &MCLEAN 1990, BOUCHET &ROCROI 2005, etc.). Others, though their initiatives are different, suggested either family rank again (MONARI et al. 1996, MCLEAN &KIEL 2007, both within Turbinoidea) or only a tribe level (GRÜNDEL 2008, within Ataphridae, Turbinoidea). The available information about shell structure of the living genera, added to Collo- niidae, shows inconsistency. Differently from COSSMANN (1918, lack of nacre), HICKMANN &MCLEAN (1990) states that “interior varies from having well-developed nacre to no visible nacre” in Colloniinae genera. The same statement is repeated also in MCLEAN &KIEL (2007) revised Colloniidae diagnosis, besides expressing of outstanding importance of the operculum morphology. This would mean either that such shell microstructure differences have in reality low/no diagnostic value, or the key characters from the morphology and life position of the opercula are less suitable for classification. Resolution of this problem is not aimed now because the original shell structures have been seldom preserved and the opercula in life position are even rarer in the fossil specimens so these diagnostic methods are mostly not applicable. To support the practice, the family needs a determination, which

Annls hist.-nat. Mus. natn. hung. 104, 2012 422 J. Szabó is based on the remaining shell characters; the keys to this attempt are in the original Collo- niidae interpretation (COSSMANN & PEYROT 1917, COSSMANN 1918). As in Collonia, a marked, single spiral morphological element of the basal shell parts develops in many, more or less turbiniform gastropod species, which have also or lack a phaneromphalus either only in adult or in all growth stages. On the basis of these characters, a considerable number of long known and recently recognised Jurassic and Triassic genera seem to be also related or belonging to Colloniidae: Lewisiella STOLICZKA, 1868, Trochotectus CONTI et FISCHER, 1984, Triangaphrus GRÜNDEL, 2003, Angulataphrus GRÜNDEL, 2003, Moreanellus FISCHER et WEBER, 1997, Bakonyia SZABÓ, 1981, Vertesella gen. n. and some still unpublished forms. The former assignments of these genera outline also a probable subfamily subdivision: Colloniinae, Lewisiellinae GRÜNDEL, 2008 and Moreanellinae FISCHER et WEBER, 1997. COSSMANN (1918) thought of Colloniidae as possibly related to the Silurian Craspe- dostoma LINDSTRÖM, 1884 and members of Ataphridae, respectively. On shells of transi- tional morphologies, SZABÓ et al. (1993) concluded also probable phylogenetic relations with ataphrids. Therefore MONARI et al. (1996) expanded Colloniidae COSSMANN, 1917 with inclusion of the former Ataphridae and Crossostomatidae as subfamilies, and added the newly recognised Adeorbisininae. In this unification, Colloniidae seemed to be the oldest available name against “Ataphridae COSSMANN, 1918” and Crossostomatidae COX in KNIGHT et al., 1960. However, recently BOUCHET &ROCROI (2005) realized that COSSMANN’s (1918) declaration about Ataphridae as “new family”, adopted into most subsequent classifications, does not correspond to the real date of establishing. This taxon was first outlined as a subfamily amongst the remarks in description of a new genus (Aulaco- trochus) already in COSSMANN’s (1915) paper. The correct family name of the taxa, unified in accordance with MONARI et al. (1996), should have been Ataphridae COSSMANN, 1915 as it is used in GRÜNDEL’s (2008) classification. Substantially, MONARI et al. (1996) and GRÜNDEL (2008) built their classification proposals on similar empirical criteria. There is an abundant and rapidly increasing group of Mesozoic (mainly Jurassic) genera that have similar, simple shell morphology, and their differences appear mainly in the peristome characters, but the former classifications have not provided appropriate frame to reflect their diversity and the phylogenetic connections. The main difference between the last two classifications is in the suggested systematic place for Proconulinae COX in KNIGHT et al., 1960 (Trochidae). MONARI et al. (1996) did not remove Proconulinae from the trochid relation, but GRÜNDEL (2000) reinterpreted this subfamily as Proconulidae while decreased the number of genera belonging to, and then placed them near to Ataphridae as sister group (GRÜNDEL 2008) within Turbinoidea. Without doubt, a few Ataphrus species of higher spire and (sub)angular periphery have shells similar to Proconulus. The species of the latter genus are ornamented already in early growth phases at least spirally but ataphrids are mostly smooth; only fine, obscure spiral threads may appear exceptionally in Ataphrus, the only genus from Ataphridae, which has comparable peristome construction to Proconulus.

Annls hist.-nat. Mus. natn. hung. 104, 2012 Vertesella, a new genus in a Middle Jurassic gastropod faunula (Colloinidae) 423

Ataphroidea COSSMANN, 1915 (nom. transl. herein ex Ataphridae COSSMANN, 1915)

Synonym – Colloniidae in sense of MONARI et al. (1996).

Diagnosis – Small to medium-sized shells (few mm to few cm) of turbiniform, conoidal to (sub)lenticular shape. Mostly thick-walled shells with obtuse apex, cyrtoconoidal spire outline and flat to convex whorls; convexity may change into single angulation or strengthen as sharp carina of various length. Protoconch depressed or planispiral, usually smooth, but single angulation and few spiral threads also found on first whorl. Periphery rounded, rarely angular or carinate. Abapical and/or abaxial deviation from normal coiling may appear in last whorl. Base convex, anomphalous, falsomphalous, cryptomphalous or phaneromphalous. Single, more or less prominent spiral carina, cord, thread or angulation may encircle central part of base with marked peristomal end. Peristome most commonly clearly continuous, rarely poorly developed at parietal region; usually simple or sometimes thickened externally and/or internally. Callosity along inner lip frequently appears. Morphology of inner lip region provides most of distinctive characters within Ataphroidea.

Remarks – Families of Ataphroidea are: Ataphridae COSSMANN, 1915 (Ataphrinae, Crossostomatinae COX in KNIGHT et al. 1960, Adeorbisinae MONARI,CONTI et SZABÓ, 1996 and Colloniidae COSSMANN, 1917 (Colloniinae, Moreanellinae FISCHER et WEBER, 1997, Lewisiellinae GRÜNDEL, 2008). These families were usually placed in Trochoidea or Turbinoidea but their relationships to the extant forms have yet been poorly defined. This may be the reason why BOUCHET &ROCROI (2005) classified Ataphridae without superfamily assignment in Vetigastropoda. Recent studies on molecular phylogeny (WILLIAMS &OZAWA 2006) suggested that extant “colloniini” are to be distinguished as Colloniidae from Turbo and closely allied genera (Turbinidae s. str.). By these results, turbinids (s. str.) are more closely related to Trochidae than to the “colloniini” and some other groups of formerly turbinid genera; non- monophyly of the calcareous opercula has become also likely. While the distance of both Ataphridae and Colloniidae from the former systematic positions increase, the shell morphological details (SZABÓ et al. 1993, MONARI et al. 1996) rather firmly suggest their close phylogenetic relation; introduction of superfamily Ata- phroidea aims to indicate this evidence. The available little shell structure information shows that Ataphroidea belongs to Vetigastropoda.

Colloniidae COSSMANN, 1917

Type genus – Collonia GRAY, 1850 (type species: Delphinula marginata LAMARCK, 1804; Eocene, Paris Basin).

Diagnosis (emended) – Moderately high to low turbiniform, rotelliform or sublenticular shells, usually with cyrtoconoidal spire outline. Aperture subcircular; peristome

Annls hist.-nat. Mus. natn. hung. 104, 2012 424 J. Szabó continuous. Protoconch depressed and having flattened to convex whorls. Outer surface of teleoconch whorls evenly arched in axial section or having one or two angulations. Base convex, anomphalous (pseudomphalous) to moderately phaneromphalous. Adaxial region of base surrounded by marked, single spiral elevation (thread, cord, keel, ridge, angulation, swollen belt). Its terminal part appears mostly as triangular or lunuliform area on outer face of inner lip at foot of columella. Operculum calcareous, and having conispiral inner and centrally pitted, flattened outer sides. Orientation of growth-lines (and peristome) slightly prosocline between sutures (or suture and periphery); basal growth-lines feebly parasigmoidal. Peri-axis elevation of base smooth or more frequently ornamented (nodes, spiral threads, fine to strong ribs). Whorls smooth or ornamented by spiral threads or striae, rarely also by collabral threads and/or riblets.

Remarks – The above characters fit COSSMANN’s (1918) COSSMANN &PEYROT’s (1917) and original descriptions, and sketches a morphological variability, abstracted from the early genus composition of the family. In dubious cases, the character state in the type species of the nominotypical genus is indicated. Presence or absence of an umbilicus and the joint occurrence of these two possibili- ties are regarded as lower than family level characters, just like the width of an umbilicus. The morphological details of the peri-axis elevation regarded as having significance for intra-family distinctions. This interpretation of Colloniidae accumulates a rather diverse group of fossil genera that needs a further, detailed taxonomic subdivision. Genera from some family group taxa, having already been recognised, as Lewisiellinae GRÜNDEL, 2008 and Moreanellinae FISCHER et WEBER, 1997, regarded as belonging to Colloniidae.

Subfamily? Colloniinae COSSMANN, 1917

gen. n.

Type species – Vertesella vertesensis sp. n.

Etymology – From Vértes Mts; vértes (Hungarian) = armoured; from lots of heavy armour pieces, thrown off by beaten medieval invaders in order to be able to escape.

Diagnosis – Sublenticular shell; juvenile whorls nearly planispiral, subsequent ones sligthly turn downwards (abapically). Initial shell part broken, however, fragment of earliest visible whorl indicate acute protoconch. Spire lower than base. Periphery becomes sharply angular then markedly carinate in last growth phase; widely rounded second angulation appears on base nearly concurrently. Moderately broad phaneromphalus (or pseudomphalus) present on base; swollen and rounded-angular rim encircles umbilicus. Peristome not preserved; broken shell part indicates thick columellar lip. Growth lines orthocline on early whorls then change into slightly prosocline and feebly prosocyrt between sutures or

Annls hist.-nat. Mus. natn. hung. 104, 2012 Vertesella, a new genus in a Middle Jurassic gastropod faunula (Colloinidae) 425 adapical suture and periphery; basal growth-lines slightly parasigmoidal in orthocline position between periphery and rim of umbilicus. Rows of short, dense riblets visible in narrow subsutural and periumbilical belts, respectively.

Remarks – The only available shell shows three different ontogenetic parts so probably represents an adult specimen in spite of its minute size. However, it may be a young individual of a larger species as well. The sublenticular shell is reminiscent to also Helicocryptus D’ORBIGNY, 1850 but this genus lacks the colloniid peri-axial elevation from the base; its shape shows closer relation to Ataphridae (Helicocryptinae COX in KNIGHT et al. 1960). Moreanellus FISCHER et WEBER, 1997 (Moreanellinae FISCHER et WEBER, 1997) has also lenticular shell with a strong peri-umbilical keel that is markedly different from the rounded-angular swollen belt around the umbilicus of Vertesella gen. n. Moreanellus has no peripheral keel The relatively large Bakonyia SZABÓ, 1981 (Colloniidae, Bajocian) has a shell of comparable early morphology. Both genera have depressed initial shells but the visible surface of the whorls are angular (carinate) on the proconch of Bakonyia then fairly convex while plane in Vertesella. Because of their spiral angulation on the whorls, Angulataphrus GRÜNDEL, 2003 and Triangaphrus GRÜNDEL, 2003 must be also compared. In Angulataphrus, the spire is rather high and the angulation occupies a midwhorl position along the whole teleoconch while it is peripheral and restricted to the last whorl in prolongation of the suture line in Vertesella that has also low spire. In Angulataphrus, the peri-umbilical swollening is not ribbed. The basal and peristomal morphology of Triangaphrus suggests also belonging to Colloniidae; a peri-umbilical swollen belt terminates in a rather large, triangular area at the foot of columella. Its whorls approach a semi-globular (strongly cyrtoconoidal) shape on the spire side but has a relatively low base. Contrariwise, Vertesella has very low spire and high base. Both genera have peripheral angulation that appears only on the last 3/4 whorl in Vertesella but it is present along the last two or three whorls in Triangaphrus; the angulation strengthens as carina only in Vertesella. Triangaphrus sp. GRÜNDEL (2003, Helicocryptinae) = Lensataphrus sp. in GRÜNDEL, NÜTZEL,KAIM &LITTLE (2011, Lewisiellinae) has also sublenticular shape with low spire and a single angulation. The non-peripheral (midwhorl) position and the length of the angulation, the lack of carina as well as the more robust peri-umbilical elevation distinguishes this problematic species from Vertesella n. gen. (but reminds the protoconch of Bakonyia). The sublenticular shells of Lensataphrus GRÜNDEL,NÜTZEL,KAIM et LITTLE, 2011 (?Lewisiellinae) have two equally developed (carinate or simple) peripheral angulations, which delimit a convex to concave outer face of the whorls in a position nearly parallel to the coiling axis. In Vertesella, first a single, carinate angulation develops peripherally on the last 3/4 whorl, then a second but hardly visible, widely rounded angulation appears (without carina) well below the former one on the base; the surface between them slightly slopes adaxially. Lensataphrus has sparse (10–12), wrinkle-like ribs that are strongest on the peri-umbilical ridge; the number of the weak riblets in similar position of Vertesella is 90–100.

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Though abstracted base morphology of Lensataphrus shows similarity to those of the colloniids, its shape vith the biangular whorls and a distinct outer face is strongly different from the ataphroidean shells, but quite near to the depressed Amphitrochus COSSMANN, 1907 species (Nododelphinulidae).

sp. n. (Figs 9–15)

Holotype – Figs 9–15; inventory number.: J.2012.1. (GGIH)

Type locality – Csóka-hegy, N from village Csókakõ, Vértes Mts, Hungary.

Type strata – Bajocian (Middle Jurassic) fissure-filling limestone (?Csókakõ Formation).

Etymology – See name of the genus (above).

Diagnosis – See diagnosis of the genus (above).

Material – One specimen; preserved diameter: 4.5 mm, height: 2 mm.

Shape – Low rotelliform-sublenticular, rather thick-walled shell of few, quite rapidly increasing whorls, which are separated by slightly impressed, groove-like suture. The earliest preserved shell part is planispiral (but protoconch lacking); its whorls are plane. How- ever, the last two whorls feebly turn abapically and their surface becomes slightly convex. A peripheral angulation develops on the last whorl from the evenly arched penultimate whorl surface nearly in continuation of the suture line. On latest parts of this angulation marked, sharp keel developed. Base is moderately phaneromphalous; a swollen shell belt borders the umbilicus with a rounded-angular adaxial margin. A third but widely rounded spiral angulation also developed medially on the base a little later than the peripheral one. Peristome is not preserved; only a surface of breakage indicates a thick umbilical lip.

Sculpture – Growth-lines are fine, their orientation is orthocline (radial) on the preserved earliest whorl then becomes slightly prosocline as a whole but feebly prosocyrt in a narrow, subsutural belt. In the same belt, packages of the neighbouring several growth- lines are fused into short riblets, just like on and near the peri-umbilical elevation. Between the periphery and the rim of the umbilicus, the growth-lines have slightly parasigmoidal shape. The shell has also remnants of a thin, shiny, outermost glaze layer that has been secreted along strongly opisthocline, faint “growth-lines”, which are almost perpendicular to the true growth-lines. Under the glaze, subregularly arraged small, dark spots are preserved probably from the original colour pattern (Figs: 14–15).

Annls hist.-nat. Mus. natn. hung. 104, 2012 Vertesella, a new genus in a Middle Jurassic gastropod faunula (Colloinidae) 427

Remarks – Protoconch has broken but remnants show it has been coaxial with the subsequent shell parts, and possibly trochospirally coiled, i.e. feebly elevated from the plane of the juvenile shell part.

Distribution – (Late ?) Bajocian.

NOTICE TO THE CSÓKA-HEGY OPERCULUM

The morphology of the calcareous operculum (Figs 5–7) corresponds quite well to that of the plan of a colloniid one (a multispiral, conoidal side and a feebly concave other side with a central pit). In Collonia, the pitted surface of the operculum has been found externally and, as the experimented rule says for dextral shell, it is sinistrally coiling. The Csóka-hegy find is constructed just inverse way; it could belong to a hypothetic, left- handed colloniid. In realty, there is no well-established correlation to any species.

CONCLUSION

The followed classification, that is a variety of proposal of MONARI et al. (1996), outlines a major cluster of the evolutionary connections, inferred from the shells of a diverse group of predominantly Mesozoic to Early Cai- nozoic (Palaeogene) gastropods. Their simple shells of lenticular, turbiniform or conoidal shape and poor ornament compose a rather homogenous morphological group (Ataphroidea). By means of this superfamily level distinction, the author intends to express that relations of these gastropods are insufficiently understood to the major groups (Trochoidea, Turbino- idea) to which they have been traditionally linked. Two internal groups of more closely related genera are distinguishable within this superfamily: Ataphridae (Ataphrinae, Crossostomatinae, Heli- cocryptinae and Adeorbisininae) and Colloniidae (Colloniinae, Lewisielli- nae, Moreanellinae). Representatives or homoeomorphs of both main shell plans seem to be sparsely present already in some Triassic faunas but their high(est) frequency and diversity have been observed since the Early Juras- sic. Ataphridae have seemed to be lacking from post-Palaeogene strata but BOUCHET &ROCROI (2005) suggest Trochaclididae as synonymous to Ataphridae on WARÉN’s unpublished data. That would mean the family has also living representatives.

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Most classifications, based on extant and/or young Cainozoic material, consider Homalopomatinae, Botropomatinae and Petropomatinae as related or synonymous to Colloniinae (and to Turbinidae). Shell morphology of their genera rarely supports these relations. Therefore, an important taxonomical-methodological question arises: is the application of the family name “Colloniidae” right or not to extant forms at all? However, to trace the latest evolution of Colloniidae is beyond the scope of this paper.

Acknowledgements – The author is grateful to STEFANO MONARI (University of Pa- dova) for critical reading of the manuscript and for his helpful suggestions.

REFERENCES

BOUCHET PH. & ROCROI J.-P. 2005: Classification and Nomenclator of gastropod families. – Malacologia (1–2): 1–397. BUDAI T., CSÁSZÁR G., CSILLAG G., FODOR L., GÁL N., KERCSMÁR ZS., KORDOS, L., PÁLFALVY S. & SELMECZI I. 2008: A Vértes-hegység földtana. (Geology of the Vértes Hills.) – Geological Institute of Hungary, Budapest, 368 pp. CONTI M. A. & FISCHER J.-C. 1983: Revisione della Fauna Mesogiurassica di Acque Fredde (Lago di Garda) Descritta da Parona 1894. – Bollettino del Museo Civico di Storia Naturale : 489–522. CONTI M. A. & FISCHER J.-C. 1984: La Faune a Gastropodes du Juarassique Moyen de Case Canepine (Umbria, Italie), Systématique, Paléobiogéographie, Paléoécologie. – Geologica Romana : 125–183. CONTI M. A. & SZABÓ J. 1987: Comparison of Bajocian gastropod faunas from the Bakony Mts (Hungary) and Umbria (Italy). – Annales historico-naturales Musei nationalis hungarici : 43–59. COSSMANN M. 1915: Étude complementaire sur le Charmoutien de la Vendée. – Mémoires de la Société linnéenne de Normandie, Section geologique : 113–159. COSSMANN M. 1918: Essais de paléontologie comparée, Vol. 11. – M. Cossmann, Paris, 388 pp. COSSMANN M. & PEYROT A. 1917: Conchologie néogénique de l’Aquitaine. Scapholodes et Gastropodes. – Actes de la Société linnéenne de Bordeaux : 157–365. FÜLÖP J., HÁMOR G., HETÉNYI R. & VÍGH G. 1960: A Vértes-hegység jura idõszaki kép- zõdményei. (Über die Jurabildungen des Vértesgebirges.) – Földtani Közlöny (1): 15–26. FÜLÖP J. 1971: Les formations jurassiques de la Hongrie. – Annales Instituti Geologici Publici Hungarici (2): 31–62.

Annls hist.-nat. Mus. natn. hung. 104, 2012 Vertesella, a new genus in a Middle Jurassic gastropod faunula (Colloinidae) 429

GALÁCZ A. 1976: Bajocian (Middle Jurassic) sections from the Northern Bakony (Hunga- ry). – Annales Universitatis Scientarium Budapestensis de Rolando Eötvös nominatae, Geologica : 177–191. GALÁCZ A. 1995: Revision of the Middle Jurassic ammonite fauna from Csóka-hegy, Vértes Hills (Transdanubian Hungary). – Hantkeniana : 119–129. GRÜNDEL J. 2000: Archaeogastropoda aus dem Dogger Norddeutschlands und des nord- westlichen Polens. – Berliner geowissenschaftliche Abhandlungen, E : 205–253. GRÜNDEL J. 2003: Gastropoden aus dem Unteren Lias (Ober-Hettangium bis Unter-Sine- murium) Südwestdeutschlands. – Stuttgarter Beiträge zur Naturkunde, B : 1–55. GRÜNDEL J. 2008: Remarks to the classification and phylogeny of the Ataphridae Cossmann, 1915 (Gastropoda, Archaeogastropoda) in the Jurassic. – Neues Jahbuch für Geologie und Paläontologie, Abhandlungen (2): 177–199. GRÜNDEL J., NÜTZEL A., KAIM A. & LITTLE C. T. S. 2011: Early Jurassic gastropods from England. – Palaeontology (3): 481–510. HICKMANN C. S. & MCLEAN J. H. 1990: Systematic revision and suprageneric classification of trochacean gastropods. – Natural History Museum of Los Angeles County Science Series, : 1–169 KNIGHT J. B., COX L. R., KEEN A. M., SMITH A. G., BATTEN R. L., YOCHELSON E. L., LUDBROOK N. H., ROBERTSON R., YONGE C. M. & MOORE R. C. 1960: Mollusca – General features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda – Gene- ral features, Archaeogastropoda and some (mainly Paleozoic) Caenogastrpoda and Opisthobranchia. – In: MOORE R. C. & PITRAT C. W. (eds): Treatise on Invertebrate Paleontology, Part I, Mollusca 1. The University of Kansas Press, Lawrence, XXIII+351 pp. KONDA J. 1970: A bakony-hegységi jura idõszaki képzõdmények üledékföldtani vizsgálata. (Litologische und Fazies-Untersuchungen der Jura-Ablagerungen des Bakony-Gebirges.) – Annales Instituti Geologici publici Hungarici, (2): 1–260 . MCLEAN J. & KIEL S. 2007: Cretaceous and living Colloniidae of the redefined subfamily Petropomatiinae, with two new genera and one new species, with notes on opercular evolution in turbinoideans, and the fossil record of Liotiidae (Vetigastropoda: Tur- binoidea). – Paläontologische Zeitschrift (3): 254–266. MONARI S., CONTI M. A. & SZABÓ J. 1996: Evolutionary systematics of Jurassic Trocho- idea: the family Colloniidae and the subfamily Proconulinae. – In: TAYLOR, J. (ed.): Origin and evolutionary radiation of the Mollusca. Oxford University Press, Oxford, pp. 199–204. PALLINI G. 1982: Una fauna a piccole ammoniti del Bajociano inferiore dell’Umbria Meri- dionale. – Quaderni di Paleontologia Stratigrafica ed Evoluzione : 129–130. PCHELINCEV V. F. & KOROBKOV I. A. 1960: Molljuski–Brjuchonogie. – In: ORLOV,Y. (ed.): Osnovy paleontologii. Izdatelstvo Akademii Nauk, Moscow, 360 pp. STURANI C. 1971: Ammonites and stratigraphy of the «Posidonia alpina» beds of the Venetian Alps. – Memorie degli Istituti di Geologia e Mineralogia dell’ Università di Padova : 1–190.

Annls hist.-nat. Mus. natn. hung. 104, 2012 430 J. Szabó

SZABÓ J. 1981: Lower and Middle Jurassic Gastropods from the Bakony Mountains (Hun- gary). Part III. Patellacea and Trochacea (Archaeogastropoda). – Annales historico- naturales Musei nationalis hungarici : 55–67. SZABÓ J. 1983: Lower and Middle Jurassic Gastropods from the Bakony Mountains (Hun- gary). Part V. Supplement to Archaeogastropoda; Caenogastropoda. – Annales historico-naturales Musei nationalis hungarici : 27–46. SZABÓ J., CONTI M. A. & MONARI S. 1993: Jurassic gastropods from Sicily; new data to the classification of Ataphridae (Trochoidea). – Scripta Geologica (Special Issue) : 406–416. VÍGH G. 1968: Jura idõszaki képzõdmények (Jurassic fomations). – In: SZENTES F. (ed): Magyarázó Magyarország 200 000-es földtani térképsorozatához. [Explanatory to 1:200 000 series of geological maps of Hungary.] L-34-I Tatabánya. Magyar Állami Földtani Intézet, Budapest, pp. 29–41. VÖRÖS A. & DULAI A. 2007: Jurassic brachiopods of the Transdanubian Range (Hungary); stratigraphical distribution and diversity changes. – Fragmenta Palaeontologica Hun- garica : 51–68. WENZ W. 1938–44: Gastropoda. Teil 1: Allgemeiner Teil und Prosobranchia. – In: SCHINDEWOLF O. H. (ed.): Handbuch der Paläozoologie. Band 6. Verlag Gebrüder Born- träger, Berlin, 1639 pp. WILLIAMS S. T. & OZAWA T. 2006: Molecular phylogeny suggests polyphyly of both the turban shells (family Turbinidae) and the superfamily Trochoidea (Mollusca: Veti- gastropoda). – Molecular Phylogenetics and Evolution : 33–51.

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