Potamididae H. Adams & A. Adams, 1854

Harzhauser, Mathias, Guzhov, Aleksandr & Landau, Bernard, 2023, A revision and nomenclator of the Cainozoic mudwhelks (Mollusca: Caenogastropoda: Batillariidae, Potamididae) of the Paratethys Sea (Europe, Asia), Zootaxa 5272 (1), pp. 1-241 : 7-59

Family Potamididae H. Adams & A. Adams, 1854

Genus Potamides Brongniart, 1810

Type species. Potamides lamarckii Brongniart, 1810 ; by monotypy ( Brongniart 1810: 368). Oligocene, France.

Note. Kadolsky (1995: 15) assumed that the original spelling lamarkii is based on the latinized name Lamarkius for Lamarck and should not be emended to lamarcki . Pacaud (2019), in contrast, pointed out that Brongniart (1810) introduced the species as “ Potamides de Lamarck ” and therefore, he considered lamarkii to be a lapsus calami, which should be corrected according to ICZN 1999, Article 32.5.1. Herein, we follow the latter view.

Illustrated material of Potamides lamarckii Brongniart, 1810 . MNHN.F.A42193, SL: 27.0 mm, MD: 9.0 mm, Ormoy-la-Rivière ( France), Rupelian, Figs 3A 1 –A 3; MNHN.F.A49861, SL: 28.3 mm, MD: 9.0 mm, Ormoy-la-Rivière ( France), Rupelian, Figs 3B 1 –B 2.

Original diagnosis. “ Taille moyenne, généralement peu élevée; forme turriculée, assez étroite; spire non étagée, à galbe conique; tours nombreux, peu élevés, à sutures profundes, plans ou convexes, ornés de rangées spirales de granulations, souvent avec une crète médiane, et de costules d’accroissement sinueuses ou incurvées, don’t quelques-unes, notamment sur le dernier tour, deviennent sublamelleuses, sans former cependant de varices régulières, mais indiquant des arrèts de l’accroissement de l’ouverture. Dernier toua peine supérieur au quart de la hauteur totale, arrondi à la péripherie de la base sur laqulle persistent les cordons granulaux et les plis sinueux jusque sur le cou qui Presque nul. Ouverture petite, arrondie dans son ensemble, avex une très faible gouttière postérieur, non limitée par unse côte pariétale; en avant et à droite, l’ouverture est échancrée, plutôt que véritablement canaliculée, […] labre peu épais, lisse et verni à l’intérieur, excavé latérlaement, […]; columelle très courte, oblique, calleuse, lisse et excavée […]; bord columellaitre étroit quoique épais, bien appliqué sur la base, sans aucune fente ombilicale.” [Size medium, generally not very high; turriculate, rather narrow; spire not gradate, conical; whorls numerous, not very high, with deep sutures, straight-sided or convex, decorated with granulated spiral cords, often with a median keel, and sinuous or curved axial ribs, of which some, in particular on the last whorl, become sublamellar, without forming, however, regular varices, but indicating growth cessations. Last whorl about one quarter of the total height, rounded at periphery of the base on which the granular cords and the sinuous ribs persist to the neck. Aperture small, circular, with a very weak anal canal, not limited by a parietal denticle; in front and on the right, the aperture is indented, rather than truly canaliculated, […]; outer lip not very thick, smooth and glossy inside, laterally excavated, […]; columella very short, oblique, callus, smooth and excavated […]; inner lip narrow but thick, well attached to base, without umbilical slit.] ( Cossmann 1906: 103).

In addition, Alimen (1948) documents a turreted protoconch of about three convex, smooth whorls (but see discussion below), early teleoconch whorls with two spiral cords, becoming beaded on second teleoconch whorls and a third spiral cord intercalated adapically on 5 th teleoconch whorl.

Stratigraphy and paleogeography. Potamides first appeared during the middle Eocene with species, such as Potamides lapidorum ( Lamarck, 1804) and Potamides perditus Bayan, 1870 , but many species listed by Le Renard & Pacaud (1995) from the Eocene of the Paris Basin as Potamides sensu stricto might need revision. The genus was well established during the Oligocene in the Northeastern Atlantic and Proto-Mediterranean Sea and entered the Central Paratethys during the Early Miocene. Potamides vanished from the Central Paratethys with the onset of the Pannonian (Tortonian) but persisted in the Eastern Paratethys from the Bessarabian to the Maeotian (Late Miocene) and appeared again in the Akchagylian (Late Pliocene/Pleistocene). In the Mediterranean Sea, the genus persisted into the Middle Pleistocene ( Brunetti 2013). To our knowledge, Potamides did not enter the Indo-West Pacific Region.

Ecology. Paratethyan Potamides species are documented from various coastal marine settings, ranging from brackish water environments to hypersaline shoals ( Harzhauser & Kowalke 2002; Latal et al. 2004). The genus, however, was not typical for mangrove environments and swamps, where Mesohalina , Ptychopotamides and Terebralia predominated.

Discussion. There was a long controversy about the use of Potamides or Pirenella for the Cainozoic European species (e.g., Kadolsky 1984; Lozouet 1986; Kowalke 2001). Reid et al. (2008) and Reid & Ozawa (2016), however, concluded that Pirenella originated in the Indo-West Pacific Region (e.g., Burdigalian of Sri Lanka in Reuter et al. 2021) and invaded the Mediterranean Sea subsequently during the Late Neogene. Such dispersal does not necessarily require a marine connection because potamidids are prone to dispersal by migratory birds ( Plaziat 1993; Kowalke 2001). Therefore, the modern distribution of Pirenella conica in the Mediterranean Sea is not related to the fossil history of Potamides .

The genus has been described as comprising species with three protoconch whorls, but species with paucispiral protoconchs occur as well (e.g., Potamides disjunctus ). As with numerous other gastropod genera, protoconch type is not a reliable generic character.

Synonyms

Genus Inderia Kolesnikov, 1950

Type species. Potamides caspius Andrussow, 1902 , original designation by Kolesnikov (1950: 96); Late Pliocene/ Pleistocene, Asia.

Discussion. Kolesnikov (1950) provided a very brief description and diagnosis, stating only: “ small, narrow shell with evenly increasing (10–11) whorls. The channel is wide, imperceptible passing into the margins of the aperture, sometimes almost disappearing. … This subgenus differs from Potamides in its very wide, sometimes disappearing canal “ [translated from Russian]. Kolesnikov(1950) might have been misguided by the rather fragmentary preservation of the specimens illustrated by Andrussow (1902). Fully preserved specimens, such as those illustrated by Ali-Zade (1967, pl. 46), show a siphonal canal, which is well demarcated from the inner and outer lips. Therefore, we see no feature that would allow to separate the type species of Inderia from Potamides . Therefore, Inderia Kolesnikov, 1950 is considered a subjective junior synonym of Potamides Bruguière, 1792 herein.

Genus Bogatschevia A.A. Ali-Zade & Kabakova in Ali-Zade, 1969

Type species: Bogatschevia bogatschevi , original designation by Ali-Zade & Kabakova in Ali-Zade (1969: 155); Late Pliocene/Pleistocene, Azerbaijan .

Discussion. The brief diagnosis just states that the new subgenus “ differs from known Akchagylian Cerithiidae by a shell of few whorls, forming a low cone, separated by a deep suture.” [translated from Russian]. This genus is based on two specimens, of which only one was illustrated by Ali-Zade & Kabakova in Ali-Zade (1969) as Bogatschevia bogatschevi . The stout shell has few extraordinarily swollen and depressed whorls, which makes it unique within all potamidids. Nevertheless, it is highly unlikely that a species of a group that usually comprises gregarious species living in large populations, occurs in such low numbers in a restricted environment with highly endemic fauna. Therefore, we assume that this genus is based on one or two highly aberrant specimens of Potamides caspius . Examples for genera that were based on aberrant or pathologic specimens of widespread potamidid species are Djeddilia Jousseaume, 1894 (= Pirenella Gray, 1847 ) and Escoffieria Fontannes, 1880 (= Potamides Bruguière, 1792 ) (see Lozouet 2003).

Genus Ashrafiella Iljina in Nevesskaja, Goncharova, Iljina, Paramonova, Popov, Babak, Bagdasaryan & Voronina, 1986

Ashrafiella nov. nom. —Iljina in Nevesskaja et al. 1986: 95.

Discussion. Ashrafiella was introduced by Iljina in Nevesskaja et al. (1986: 95) as new name for Bogatschevia Ali-Zade & Kabakova in Ali-Zade, 1969 non Jatzko, 1962 [ Unionidae ]. Jatzko (1962), however, did not designate a type species for his new genus, which he did in Jatzko (1972). Therefore, Ashrafiella is an objective junior synonym of Bogatschevia Ali-Zade & Kabakova in Ali-Zade, 1969. Bogatschevia Jatzko, 1972 , in contrast will need a new name. Note that Nevesskaja et al. (1986: 5) stated that L.B. Iljina was the author of the gastropod taxa in that monograph.

Species-level taxa placed in Potamides reported from the Paratethys aktschagylicum . Cerithium . A.A. Ali-Zade, 1967 → Potamides caspius Andrussow, 1902 bellum . Cerithium . A.A. Ali-Zade & Petrova in A.A. Ali-Zade, 1969

→ Potamides caspius Andrussow, 1902 bigranulata. Cerithium . Karlov, 1932 → Potamides taitboutii (d’Orbigny, 1844) bogatschevi . Bogatschevia . A.A. Ali-Zade & Kabakova in A.A. Ali-Zade, 1969

→ Potamides caspius Andrussow, 1902 brenneri . Cerithium . Handmann, 1883b → Potamides fraterculus (Mayer, 1878) carum . Cerithium . A.A. Ali-Zade & Petrova in A.A. Ali-Zade, 1969 → Potamides caspius Andrussow, 1902 caspius . Potamides . Andrussow, 1902 → Potamides caspius Andrussow, 1902 comroicus . Potamides . K.A. Ali-Zade, 1940 → Potamides caspius Andrussow, 1902 conica . Cerithium . Simionescu & Barbu, 1940 → Potamides disjunctus (J. de C. Sowerby, 1832) connexum . Cerithium . Eichwald, 1851 → Potamides disjunctus (J. de C. Sowerby, 1832) constantiae . Cerithium . Stefanescu, 1897 → Potamides taitboutii (d’Orbigny, 1844) cordatum . Cerithium A.A. Ali-Zade & Kabakova in A.A. Ali-Zade, 1969

→ Potamides caspius Andrussow, 1902 coronatum . Cerithium . auctores → Potamides disjunctus ( Sowerby, 1832) danatense . Cerithium . A.A. Ali-Zade, 1967 → Potamides caspius Andrussow, 1902 deformatus . Potamides . K.A. Ali-Zade, 1940 → Potamides caspius Andrussow, 1902 disjunctoides . Cerithium . Sinzow, 1897 → Potamides taitboutii (d’Orbigny, 1844) disjunctum . Cerithium . J. de C. Sowerby, 1832 → Potamides disjunctus ( Sowerby, 1832) dobrogense . Cerithium . Simionescu & Barbu, 1940 → Potamides disjunctus (J. de C. Sowerby, 1832) dominici. Potamides (Bittium) . Hilber, 1891 → Potamides fraterculus (Mayer, 1878) ebersini . Cerithium . A.A. Ali-Zade, 1967 → Potamides caspius Andrussow, 1902 eldaricus . Potamides . Kolesnikov, 1950 → Potamides caspius Andrussow, 1902 elegantus . Potamides . Barg, 1966 → Potamides taitboutii (d’Orbigny, 1844) enodis. Potamides . Ionesi, 1968 [nomen nudum] → Potamides disjunctus (J. de C. Sowerby, 1832) extortus. Potamides . Kókay, 1954 → Potamides fraterculus (Mayer, 1878) fraterculus . Cerithium . Mayer, 1878b → Potamides fraterculus (Mayer, 1878) friedbergi . Cerithium . Simionescu & Barbu, 1940 → Potamides disjunctus (J. de C. Sowerby, 1832) galidzgensis . Pirenella . Iljina in Iljina et al., 1976 → Potamides galidzgensis (Iljina in Iljina et al., 1976) granuliferum . Bittium . auctores → Potamides sp. hartbergensis . Potamides . Hilber, 1891 → Potamides fraterculus (Mayer, 1878) hersonica . Potamides . Barg, 1966 → Potamides taitboutii (d’Orbigny, 1844) hoellesense . Cerithium . Handmann, 1883b → Potamides fraterculus (Mayer, 1878) istritzense . Cerithium . Teisseyre, 1908 → Potamides taitboutii (d’Orbigny, 1844) jagorlicus . Potamides . Barg, 1966 → Potamides taitboutii (d’Orbigny, 1844) jumudicum . Cerithium . A.A. Ali-Zade, 1967 → Potamides caspius Andrussow, 1902