The Clavatulidae (Gastropoda, Conoidea) of the Miocene Paratethys Sea with considerations on fossil and extant Clavatulidae genera
Authors: HARZHAUSER MATHIAS , LANDAU BERNARD , JANSSEN RONALD 
Published: March 2022

We present a revision of the Clavatulidae gastropods of the Neogene of the Central and Eastern Paratethys seas. In total, 111 species level names have been used in the literature for Clavatulidae of the Paratethys Sea. After revision, we document 83 species. Seventeen genus-level groups are recognized, of which eleven are formally described. Six species-groups cannot be clearly placed in a certain Clavatulidae genus. The presence of Clavatula, Perrona, Pusionella, Scaevatula and Tomellana in the Paratethyan assemblages documents a biogeographic relationship with extant Clavatulidae faunas from the tropical eastern Atlantic. No Indo-West Pacific influences were observed, as Paratethyan species previously placed in Turricula, which belongs within the Clavatulidae, do not belong within that genus. The Langhian (middle Miocene) diversity of 62 species is comparable to the number of extant species recorded from West Africa (~65 species), but displays a much higher diversity at genus level. This high biodiversity in Paratethyan assemblages suggests that the Central Paratethys was a major center of radiation for clavatulid gastropods, which is also expressed by a high endemicity of 98.8%. In contrast, the clavatulid diversity in the Eastern Paratethys was very low and stratigraphically restricted to the early Miocene Sakaraulian.
Granulatocincta nov. gen., Megaclavatula nov. gen., Neoperrona nov. gen., Olegia nov. gen., Striopusionella nov. gen. are established as new genera. Clavatula sorini nov. sp., ‘Clavatula’ irisae nov. sp., Tomellana dulaii nov. sp., Tomellana aueri nov. sp., Granulatocincta callim nov. sp., Granulatocincta theoderichi nov. sp., Megaclavatula grunerti nov. sp., Megaclavatula pilleri nov. sp., Neoperrona zoltanorum nov. sp., Olegia mandici nov. sp., Perrona koeberli nov. sp., Perrona loetschi nov. sp., Pusionella hofmanni nov. sp. are described as new species from the Miocene Paratethys, and ‘Clavatula’ ariejansseni nov. sp., ‘Clavatula’ atatuerki nov. sp. and Granulatocincta pelliscrocodili nov. sp., are described from the eastern Proto-Mediterranean Karaman Basin of Turkey.
‘Clavatula’ jarzynkae nov. nom., Perrona grossi nov. nom., Perrona ilonae nov. nom. and Perrona wanzenboecki nov. nom. are introduced as new names for Clavatula auingeri Finlay, 1927 [non Hilber, 1879], Pleurotoma (Clavatula) auingeri Hilber, 1879 [non Hoernes, 1875], Clavatula vindobonensis nodosa Csepreghy-Meznerics, 1954 [non Bellardi, 1847] and Pleurotoma concinna Handmann, 1883 [non Scacchi, 1836] respectively.
Clavatula kowalewskii Bałuk, 2003, Clavatula letkésensis Csepreghy-Meznerics, 1953, Pleurotoma aculeatum Eichwald, 1830, Pleurotoma subscalaris Handmann, 1882 and Pleurotoma (Clavatula) reginae Hoernes & Auinger, 1891 are treated as junior subjective synonyms of Pleurotoma (Clavatula) antoniae Hoernes & Auinger, 1891, Perrona emmae (Hoernes & Auinger, 1879), Pleurotoma laevigata Eichwald, 1830, Pleurotoma schreibersi Hörnes, 1854 and Pleurotoma (Clavatula) apolloniae Hoernes & Auinger, 1891 respectively.