Perrona descendens ( Hilber, 1879 )

Harzhauser, Mathias, Landau, Bernard & Janssen, Ronald, 2022, The Clavatulidae (Gastropoda, Conoidea) of the Miocene Paratethys Sea with considerations on fossil and extant Clavatulidae genera, Zootaxa 5123 (1), pp. 1-172 : 103-104

Perrona descendens ( Hilber, 1879)

Figs 31A 1 –A 3 , B 1 –B 3 , C 1 –C 3 , D 1 –D 3 , 4M , 5 , 7

Pleurotoma Jouanneti [sic] Desm. — Hörnes 1848: 20 (partim) [non Tomellana jouannetii ( Des Moulins, 1842) ]. Pleurotoma Jouanneti [sic] Des Moul.— Hörnes 1854: 346 (partim), non figs [non Tomellana jouannetii ( Des Moulins,

1842)]. * Pleurotoma (Clavatula) descendens Hilb. — Hilber 1879: 19, pl. 3, figs 5a–b. Pleurotoma (Clavatula) descendens Hilb. — Hoernes & Auinger 1891 (partim): 355, pl. 48, figs 9a–c [non figs 7–8 = Perrona loetschi nov. sp.]. Pleurotoma cf. descendens Hilb. — Bauer 1900: 25, pl. 1, figs 4a–b. Pleurotoma — Bauer 1900: 26, pl. 1, figs 6a–c. Clavatula Jouanneti [sic] Desm.— Friedberg 1912: 201, pl. 13, fig. 1 [non Tomellana jouannetii ( Des Moulins, 1842) ]. Clavatula Jouanneti [sic] Desm. var. descendens Hilb. — Friedberg 1938: 144. Clavatula (Perrona) jouanneti [sic] descendens Hilber — Csepreghy-Meznerics 1954: 53. P [errona]. (P [errona]) descendens (Hilb.) — Sieber 1958a: 157. Pleurotoma (Clavatula) veronicae R. Hoernes et Auinger — Florei 1961: 679, pl. 6, fig. 32 [non ‘ Clavatula ’ veronicae ( Hoernes & Auinger, 1891)]. Clavatula vindobonensis (Partsch) — Kecskemétiné Körmendy 1962: 92, 11, figs 2 [non Perrona vindobonensis (Quenstedt, 1884)].? Clavatula (Perrona) jouanneti [sic] (Des Moulins)— Răileanu & Negulescu 1964: 179, pl. 16, fig. 2 [non Tomellana jouannetii ( Des Moulins, 1842)]. Clavatula jouanneti [sic] descendens Hilber, 1879 — Strausz 1966: 401, pl. 14, figs 15–16. Clavatula (Perrona) descendens Hilb. — Kókay 1966: figure caption, pl. 9, fig. 19. Clavatula (Perrona) jouannetti [sic] descendens Hilber — Csepreghy-Meznerics 1972: 32, pl. 16, figs 2–3. Clavatula (Perrona) jouanettii [sic] descendens (Hilber) 1879 — Bohn-Havas 1973: 1063, pl. 6, fig. 10. Perrona (Perrona) vindobonensis nodosa (Csepreghy-Meznerics) — Nikolov 1994: 59, pl. 6, figs 14–15 [non ‘Clavatula’ nodosa

( Bellardi, 1847), nec Csepreghy-Meznerics, 1954]. Perrona carinifera ( Grateloup, 1832) — Mikuž 1998: 78, pl. 5, figs 4–5 [non ‘Clavatula’ carinifera ( Grateloup, 1832) ]. Clavatula (Perrona) jouanneti [sic] ( Des Moulins, 1842)— Mikuž 1998; 78, pl. 5, fig. 8 [non Tomellana jouannetii ( Des Moulins, 1842)]. Perrona cf. descendens ( Hilber, 1879) — Mikuž 1998: 79, pl. 5, fig. 9. Perrona (Perrona) jouanneti [sic] descendens (Hilber) — Schultz 1998: 74, pl. 30, fig. 4. Clavatula jouanneti [sic] Desmoulins— Popa & Chira 1999: 400, pl. 2, fig. 7 [non Tomellana jouannetii ( Des Moulins, 1842) ]. Perrona (Perrona) descendens ( Hilber, 1879) — Bałuk 2003: 40, pl. 9, figs 1–7. Perrona carinifera ( Grateloup, 1832) — Mikuž 2009: 33, pl. 11, fig. 148 [non ‘Clavatula’ carinifera ( Grateloup, 1832) ]. Perrona cf. descendens ( Hilber, 1879) — Mikuž 2009: 34, pl. 11, fig. 150. Perrona descendens ( Hilber, 1879) — Landau et al. 2013: 290, pl. 49, figs 8–9, pl. 50, fig. 13. Perrona descendens ( Hilber, 1879) — Kovács & Vicián 2021: 145, pl. 3, figs 11–14. Perrona sp. —Jovanović et al. 2021: 327, figs 10/9–10. non Clavatula (Perrona) jouanneti jouanneti [sic] (Des Moulins)— Švagrovský 1958: 19, pl. 6, figs 1–2 [= Perrona lydiae ( Hoernes & Auinger, 1891)].

Type material. Lectotype designated herein: NHMW 1862 /0033/0034, SL: 28.2 mm, MD: 10.5 mm, Pöls ( Austria), illustrated in Hilber (1879, pl. 3, figs 5a–b), figs 31A 1 –A 3 . Paralectotypes: NHMW 1861 /0001/0260a, SL: 29.7 mm , MD: 11.3 mm, Pöls ( Austria), figs 31C 1 –C 3 ; NHMW 1861 /0001/0260b, SL: 27.9 mm , MD: 10.7 mm, Pöls ( Austria), figs 31D 1 –D 3 .

Illustrated material. NHMW 1949 /0005/0101, SL: 26.7 mm , MD: 9.3 mm, St. Florian ( Austria), illustrated in Hoernes & Auinger (1891, pl. 48, figs 9a–c), figs 31B 1 –B 3 ; NHMW 1846 /0049/0017, 4 spec., St. Florian ( Austria) .

NHMW 1846 /0049/0017, SL: 14.0 mm . MD: 5.5 mm, St. Florian ( Austria), Fig. 4M .

Additional studied material. NHMW 1860 /0005/0044, 17 spec., Pöls ( Austria) ; NHMW 1860 /0005/0044, 15 spec., Pöls ( Austria) ; NHMW A641 , 10 spec., Bahna ( Romania) ; NHMW 1855 /0038/0033, 1 spec., Korytnica ( Poland) .

Revised description. Shell medium-sized, solid, moderately broad fusiform, with gradate spire; apical angle 36–40°. Protoconch not preserved. Teleoconch of ten whorls. Early teleoconch whorls weakly concave with two smooth, narrow, adsutural spiral cords separated by weakly concave central part with comma-shaped axial riblets. Axial riblets fading on fourth whorl. Later whorls subcylindrical with prominent, swollen, rounded subsutural collar bearing poorly defined tubercles. Delicate spiral threads on early spire whorls, subobsolete on penultimate and last whorls. Suture narrowly incised, weakly irregular. Last whorl ~60% of total height, subcylindrical. Subsutural collar broad, rounded, weakly tubercular, subsutural ramp narrow, concave, delimited by weak, rounded shoulder, weakly convex below, weakly angled at moderately constricted concave base. Siphonal fasciole weakly delimited, weakly swollen with prominent growth increments. Sculpture of weak irregular spiral cords on base, strengthening slightly over fasciole. Aperture moderately narrow, pyriform. Outer lip not thickened, smooth within. Anal sinus narrow to moderately narrow, moderately deep, asymmetrically U-shaped, with apex just below subsutural collar. Siphonal canal moderately long, moderately wide, only slightly deflected to the left with shallowly notched tip. Columella moderately excavated in upper third, straight below, weakly twisted at fasciole, smooth. Columellar and parietal callus thin, adherent, poorly delimited from base, forming broad rim.

Discussion. Perrona coppii ( Bellardi, 1877) , from the Tortonian of Stazzano ( Italy), is closely similar to P. descendens . A separation from P. descendens ( Hilber, 1879) is based on the beaded basal and peribasal spiral cords and by the presence of more prominent spiral cords on the spire whorls in P. coppii (see Ferrero Mortara et al. 1981, pl. 14, figs 8a–b).

Perrona descendens is also highly reminiscent of P. ilonae nov. nom. However, the two are immediately distinguished by the prominent spiral cords covering the entire surface below the subsutural collar in P. ilonae whereas in P. descendens spiral sculpture is restricted to the base and fasciole, and rather weak. See Perrona sabinae ( Hoernes & Auinger, 1891) for further discussions.

Paleoenvironment. Inner neritic, found from coastal mudflats to shallow sandy environments (e.g., Kleinebersdorf, Zuschin et al. 2004)

Distribution in Central Paratethys.? Eggenburgian (Early Miocene): Southern Carpathians: Sălătrucu ( Romania) ( Răileanu & Negulescu 1964); Karpatian (early Miocene): Korneuburg Basin: Weinsteig, Kleinebersdorf ( Austria) ( Hoernes & Auinger 1891); Badenian (middle Miocene): North Alpine-Carpathian Foreland Basin: Moravská Třebová (= Mährisch-Trübau) (the Czech Republic) ( Hoernes & Auinger 1891); Korytnica Basin: Korytnica ( Poland) ( Bałuk 2003); Styrian Basin: St. Florian, Pöls ( Austria) ( Hilber 1879); Oberpullendorf Basin: Ritzing ( Austria) ( Hoernes & Auinger 1891); Pannonian Basin: Letkés, Pécsszabolcs, Várpalota, Mátraverebély, Szob ( Hungary) ( Csepreghy-Meznerics 1954; Strausz 1966; Bohn-Havas 1973; Kovács & Vicián 2021); Southern Pannonian Basin: Hrvaćani ( Bosnia and Herzegovina) (Jovanović et al. 2021); Krka Basin: Ivanji dol ( Slovenia) ( Mikuž 2009); Bahna Basin: Bahna ( Romania) (hoc opus); Zorlenţul Mare Basin ( Romania) ( Florei 1961); Borod Basin: Cetea (borehole) ( Romania) ( Popa & Chira 1999); Dacian Basin: Târnene ( Bulgaria) ( Nikolov 1994).

Figures

FIGURE 1. A. Map of central and south–eastern Europe, representing the area that was partly covered by the Central Paratethys Sea. The white insert corresponds to the area that is restored in the palaeogeographic map below. B. Paleogeography of the Middle Miocene Paratethys Sea showing the most important sedimentary basins (NAFB: North Alpine Foreland Basin, E.-Sopr B: Eisenstadt–Sopron Basin).

FIGURE 3. Early teleoconch whorls of extant and fossil Clavatulidae species. A. Clavatula regia (Röding, 1798), NHMW 1683 6/28, West Africa. B. Clavatula rubrifasciata (Reeve, 1845), NHMW 10203, Gorée (Senegal). C. ‘Clavatula’ diadema (Kiener, 1839), NHMW 1884/0010/0023, Gorée (Senegal). D. ‘Clavatula’ mystica (Reeve, 1843), MNHN-IM-2013-59897, Est de l'ile de Gorée (Senegal). E. ‘Clavatula’ bimarginata (Lamarck, 1822), MNHN-IM-2013-43369, Dakar (Senegal). F. ‘Clavatula’ pyramidata (Kiener, 1839), MNHN-IM-2009-24982, Baie de Pointe-Noire (Republique du Congo). G. Clavatula sorini nov. sp., paratype, NHMW 1874/0025/0046, Lăpugiu de Sus (Romania). H. ‘Clavatula’ romana (Defrance, 1826), NHMW 1999/0098/0004, Modra-Kráľová (Slovakia). I. ‘Clavatula’ interrupta (Brocchi, 1814), NHMW 1861/0033/0420, Buituri (Romania). J. ‘Clavatula’ hirmetzli Kovács & Vicián, 2021, NHMW 1868/0001/0069, Baden-Sooss (Austria). K. ‘Clavatula’ irisae nov. sp., NHMW 2021/0126/0001, holotype, Gainfarn (Austria). L. ‘Clavatula’ sophiae (Hoernes & Auinger, 1891), lectotype, NHMW 2021/0127/0001, Enzesfeld (Austria). M. ‘Clavatula’ veronicae (Hoernes &Auinger, 1891), NHMW 1855/0043/0044b, Lăpugiu de Sus (Romania). N. ‘Clavatula’ santhai Kovács & Vicián, 2021, holotype, HNHM, PAL 2021.20.1, Letkés (Hungary).O. ‘Clavatula’ camillae (Hoernes &Auinger, 1891), lectotype, NHMW 1949/0005/0042, Lăpugiu de Sus (Romania). P. Megaclavatula amaliae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0049, Lăpugiu de Sus (Romania). Q. Megaclavatula grunerti nov. sp., paratype, NHMW 2021/0127/0003, Enzesfeld (Austria). R. Megaclavatula laevigata (Eichwald, 1830), NHMW 1853/0038/0035a, Korytnica (Poland). S. Megaclavatula pilleri nov. sp., holotype, NHMW 1949/0005/0051a, Baden-Sooss (Austria). T. Megaclavatula polonica (Pusch, 1837), NHMW 1862/0001/0324, Bad Vöslau (Austria).U. Granulatocincta angelae (Hoernes &Auinger, 1891), paralectotype, NHMW 1949/0005/0099, Möllersdorf (Austria). V. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 1846/0028/0009, Enzesfeld (Austria). W. Granulatocincta nataliae (Hoernes & Auinger, 1891), NHMW 1855/0035/0021, holotype, Belgrade/Rakovica (Serbia). X. Granulatocincta schreibersi (Hörnes, 1854), paralectotype, NHMW1997z0178/1133, Bad Vöslau (Austria).Y. Granulatocincta callim nov. sp., paratype, NHMW 2021/0128/0001, Bad Sauerbrunn (Austria). Z. Granulatocincta contorta (Švagrovský, 1958), holotype, SNMB Z 3076, Kuzmice (Slovakia), Aa. Granulatocincta theoderichi nov. sp., paratype, NHMW 2016/0026/0320b, Ritzing (Austria). Ab. Olegia doderleini (Hörnes, 1854), paralectotype, NHMW 1858/0015/0147, Wiesen (Austria). Ac. Olegia mandici nov. sp., paratype, NHMW 1872/0031/0008a, Bekića vrelo, Čremušnica at Donji Kirin (Croatia). Ae. Olegia dorotheae (Hoernes & Auinger, 1891), paralectotype, NHMW 1949/0005/0040, Kleinebersdorf (Austria). Scale bar is 5 mm for all specimens except for Granulatocincta contorta, Olegia doderleini and O. mandici.

FIGURE 4A. ‘Clavatula’ justinae (Hoernes & Auinger, 1891), NHMW ex1949/0005/0110b, Buituri (Romania). B. ‘Clavatula’ mariae (Hoernes & Auinger, 1891), lectotype, NHMW 1849/0013/0009, Mold (Austria). C. ‘Clavatula’ olgae (Hoernes & Auinger, 1891), lectotype, NHMW 1862/0001/0552, Buituri (Romania). D. ‘Clavatula’ orientoromana Báldi, 1960, holotype, M.60.145, Szokolya (Hungary). E. ‘Clavatula’ juliae (Hoernes & Auinger 1891), paralectotype, NHMW 1949/0005/0043c, Lăpugiu de Sus (Romania). F. ‘Clavatula’ apolloniae (Hoernes & Auinger, 1891), NHMW 1868/0001/0474a, Lăpugiu de Sus (Romania). G. Neoperrona harzhauseri (Kovács & Vicián, 2021), NHMW 2019/0151/019a, Letkés (Hungary). H. Neoperrona zoltanorum nov. sp., holotype, NHMW 2019/0151/0108, Letkés (Hungary). I. Scaevatula sidoniae (Hoernes & Auinger, 1891), NHMW 1855/0043/0045, holotype, Lăpugiu de Sus (Romania). J. Pusionella palatina Strausz, 1954, NHMW 2021/0159/0003, Várpalota (Hungary). K. Pusionella hofmanni nov. sp., NHMW 1867/0019/0136, holotype, Coşteiu de Sus (Romania). L. Perrona barbarae (Hoernes & Auinger, 1891) NHMW 1949/0005/0103a, Rückersdorf (Austria). M. Perrona descendens (Hilber, 1879), NHMW 1846/0049/0017, St. Florian (Austria). N. Perrona emmae (Hoernes & Auinger 1891), NHMW 2021/0127/0005, Enzesfeld (Austria), NHMW 1970/1396/1449, Baden (Austria). O. Perrona ernae (Šuklje, 1929), NHMW 2021/0156/0001, Weinsteig (Austria). P. Perrona floriana (Hilber, 1879), holotype, NHMW 1858/0020/0012, Guglitz at St. Florian (Austria). Q. Perrona ilonae nov. nom., M.61.4367, Szentkút, Mátraverebély (Hungary). R. Perrona lydiae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0112, Gainfarn (Austria). S. Perrona oliviae (Hoernes & Auinger 1891), paralectotype, NHMW 1873/0026/0027, Lăpugiu de Sus (Romania). T. Perrona rosaliae (Hoernes & Auinger 1891), lectotype, NHMW 1999z0098/0002, Vienna/Pötzleinsdorf (Austria). U. Perrona sabinae (Hoernes & Auinger 1891), paralectotype, NHMW 2021/0154/0001, Lăpugiu de Sus (Romania). V. Perrona styriaca (Hilber, 1879), lectotype, NHMW 1846/0037/0309, Pöls (Austria). W. Perrona ursulae (Hoernes & Auinger, 1891), paralectotype, NHMW 1846/0001/1294, Buituri (Romania). X. Perrona vindobonensis (Quenstedt, 1884), paralectotype, NHMW 1997z0178/1467, Gainfarn (Austria), Y. Perrona wanzenboecki nov. nom., NHMW 2021/0158/0001, neotype, St. Veit an der Triesting (Austria). Z. Tomellana dulaii nov. sp., NHMW 2021/0148/0002, holotype, Nemeşeşti (Romania). Aa. Tomellana aueri nov. sp., NHMW 2013/0078/0436, Baden (Austria). Ab. Striopusionella szekelyhidiae (Kovács & Vicián, 2021), holotype, PAL 2021.19.1, Letkés (Hungary). Ac. Scaevatula sidoniae (Hoernes & Auinger, 1891). NHMW 1855/0043/0045, holotype, Lăpugiu de Sus (Romania). Scale bar is 5 mm for all specimens except for Perrona barbarae and protoconch of Scaevatula sidoniae.

FIGURE 5. Anal sinus morphology of Paratethyan Clavatulidae.

FIGURE 7. Apertures of Paratethyan Clavatulidae (O. = Olegia, ‘C.’ = ‘Clavatula’, N. = Neoperrona, P. = Perrona, Pu. = Pusionella, Sc. = Scaevatula, S. = Striopusionella, T. = Tomellana).

FIGURE 31 A1–A3. Perrona descendens (Hilber, 1879), lectotype, NHMW 1862/0033/0034, Pöls (Austria). B1–B3. Perrona descendens (Hilber, 1879), paralectotype, NHMW 1949/0005/0101, St. Florian (Austria). C1–C3. Perrona descendens (Hilber, 1879), paralectotype, NHMW 1861/0001/0260, Pöls (Austria). D1–D3. Perrona descendens (Hilber, 1879), paralectotype, NHMW 1861/0001/0260a, Pöls (Austria). E1–E3. Perrona floriana (Hilber, 1879), holotype, NHMW 1858/0020/0012, Guglitz at St. Florian (Austria). F1–F2. Perrona ilonae nov. nom., M.61.4367, Szentkút, Mátraverebély (Hungary).