Megaclavatula amaliae ( Hoernes & Auinger, 1891 ) Harzhauser & Landau & Janssen, 2022

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 : 61-63

Megaclavatula amaliae ( Hoernes & Auinger, 1891) nov. comb.

Figs 20A 1 –A 3 , B 1 –B 3 , C 1 –C 2 , 3P , 5 , 6

Clavatula Amaliae —Hoernes 1891: 130 [nomen nudum].

* Pleurotoma (Clavatula) Amaliae nov. form.— Hoernes & Auinger 1891: 346, pl. 44, figs 1–2.

Clavatula (Clavatula) amaliae (R. Hörn. Auing.) — Sieber 1956: 245.

Clavatula (Clavatula) amaliae R. Hoernes et Auinger — Švagrovský 1958: 12, pl. 2, fig. 4

Clavatula evae ( Hoernes & Auinger, 1891) — Bałuk 2003: 34, pl. 5, figs 1–8 [non Megaclavatula evae ( Hoernes & Auinger, 1891) ].

non Clavatula laevigata Eichw. var., aff. Cl. Amaliae R. H. I A.—Friedberg 1928: 569, text-fig. 81 middle [= Megaclavatula laevigata ( Eichwald, 1830) ]. non Clavatula (Clavatula) laevigata var. amaliae (Hoernes und Auinger 1891) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 198, pl. 47, fig. 9 [= Megaclavatula laevigata ( Eichwald, 1830) ]. non Clavatula amaliae (Hoern, & Auing.) juv.— Báldi 1960: 81, pl. 3, fig. 3 [=? Megaclavatula laevigata ( Eichwald, 1830) ]. non Clavatula asperulata amaliae Hoernes & Auinger, 1891 — Strausz 1966: 407, pl. 16, figs 3–4 [= ‘ Clavatula ’ camillae ( Hoernes & Auinger, 1891)]. non Clavatula (Clavatula) amaliae (Hoernes R. & Auinger, 1891) — Mikuž, 1998: 69, pl. 1, figs 3–4 [= Megaclavatula polonica (Pusch, 1837)]. non Clavatula amaliae (R. Hoernes & Auinger, 1891) — Mikuž 2009: 30, pl. 10, fig. 132 [= Megaclavatula polonica (Pusch, 1837)]. non Clavatula amaliae ( Hoernes & Auinger, 1891) — Kovács & Vicián 2021: 141, pl. 1, figs 21–25 [= ‘ Clavatula ’ camillae ( Hoernes & Auinger, 1891)].

Type material. Lectotype designated herein: NHMW 1949 /0005/0049a, SL: 72.2 mm, MD: 24.9 mm, Lăpugiu de Sus ( Romania), illustrated in Hoernes & Auinger (1891, pl. 44, fig. 1), figs 20A 1 –A 3, 3P. Paralectotypes: NHMW 1949 /0005/0049b, SL: 69.1 mm , MD: 24.1 mm, Lăpugiu de Sus ( Romania), illustrated in Hoernes & Auinger (1891, pl. 44, fig. 2), figs 20B 1 –B 3 .

Illustrated material. NHMW 2021/0132/0001, SL: 76.6 mm, MD: 28.3 mm, Baden-Sooss ( Austria), figs 20C 1 –C 2.

Revised description. Shell large, solid, moderately broad fusiform with high, weakly coronate spire; apical angle ~35°. Protoconch not preserved. Teleoconch of at least ten whorls, distinctly concave mid-whorl. First three teleoconch whorls with tripartite spiral sculpture consisting of weak subsutural spiral cord, distinctly concave mid-whorl portion with weak cord bearing tiny beads, and prominently beaded suprasutural cord. Subsutural cord becomes beaded on fourth to fifth teleoconch whorl, forming close-set double row of beads with abapical beads of previous whorl, separated by narrowly incised, weakly undulating suture. Sculpture changing on sixth teleoconch whorl; subsutural cord becomes weak subsutural collar with small, spiny tubercles placed short distance below suture; mid-whorl cord disappears. Beads on suprasutural cord turning into short, spiny tubercles partly covered by subsequent whorl. Fine spiral threads cover entire shell surface. Last whorl 60% of total height. Poorly delimited spinous subsutural collar. Distinctly concave subsutural ramp, angled at shoulder and base. Base strongly constricted. Siphonal fasciole long, rounded, strongly twisted, with prominent growth increments, forming narrow pseudumbilical chink. Four primary spiral cords at mid-whorl, adapical delimiting shoulder, abapical delimiting base, with secondary and tertiary spirals intercalated, weaker cords over base and siphonal fasciole, one cord slightly strengthened delimiting fasciole. Prominent axial growth lines give surface finely reticulated aspect. Aperture elongate pyriform. Outer lip not thickened, smooth within. Anal sinus moderately wide and deep, broadly V-shaped, with apex below subsutural collar in deepest part of subsutural concavity. Siphonal canal long, moderately narrow, twisted, deflected to the left, shallowly notched at tip. Columella strongly excavated, twisted at fasciole, smooth. Columellar and parietal callus strongly thickened, sharply delimited.

Discussion. Megaclavatula amaliae ( Hoernes & Auinger, 1891) is a very large species documented so far only by few specimens. The somewhat misleading illustrations in Hoernes & Auinger (1891) gave rise to numerous misidentifications in the literature. All these specimens are easily distinguished from M. amaliae either by their smaller size and lower spire (e.g., Kojumdgieva in Kojumdgieva & Strachimirov 1960; Báldi 1960; Mikuž 1998, 2009) or by their slender shape and different sculpture on early whorls (e.g., Strausz 1966; Kovács & Vicián 2021). Megaclavatula amaliae resembles Megaclavatula laevigata ( Eichwald, 1830) and M. grunerti nov. sp. in size and shape, but is distinguished by its double row of beads on early teleoconch whorls, its coarse spiral cords and the reticulate sculpture at mid-whorl on the last whorl.

‘Clavatula’ heros sensu Bellardi, 1877 (non Mayer, 1858), from the Langhian or Burdigalian of the Colli Torinesi ( Italy), resembles M. amaliae (see Bellardi 1877: 182, pl. 6, fig. 5b) in size, sculpture and general shape. A clear difference is the position of the suture, which is distinctly below the shoulder of the spire whorls in the Italian species, resulting in a more elongate spire with markedly angulated whorls. The Italian species is clearly not conspecific with Clavatula heros Mayer, 1858 , described from the Langhian of Jean de Marsacq ( France), which is stouter biconic or more stoutly biconic and has the suture close below the nodes (see Mayer 1858: 81, pl. 3, fig. 6).

Paleoenvironment. The occurrences in the Baden Formation in the Vienna Basin suggest middle to outer neritic environments with up to 210 m water depth ( Kranner et al. 2021).

Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica ( Poland) ( Bałuk 2003); Vienna Basin: Baden, Bad Vöslau, Baden-Sooss ( Austria) ( Hoernes & Auinger 1891), Borský Mikuláš ( Slovakia), Mikulov (the Czech Republic) ( Švagrovský 1958); Eisenstadt-Sopron Basin: Forchtenau ( Austria) ( Sieber 1956); Făget Basin: Lăpugiu de Sus ( Romania) ( Hoernes & Auinger 1891).

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 2. Descriptive terms used in descriptions and measurements to shell morphology, whorl profiles and anal sinus. SL: shell length, MD: maximum diameter, AA: apical angle, LWH: last whorl height, AH: aperture height.AL: aperture length.AW: aperture width.

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 5. Anal sinus morphology of Paratethyan Clavatulidae.

FIGURE 6. Apertures of Paratethyan Clavatulidae (C. = Clavatula, ‘C.’ = ‘Clavatula’, G. = Granulatocincta, M. = Megaclavaltula, O. = Olegia).

FIGURE 20 A1–A3. Megaclavatula amaliae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0049a, Lăpugiu de Sus (Romania). B1–B3. Megaclavatula amaliae (Hoernes & Auinger, 1891), paralectotype, NHMW 1949/0005/0049b, Lăpugiu de Sus (Romania), C1–C2. Megaclavatula amaliae (Hoernes & Auinger, 1891), NHMW 2021/0132/0001, Baden-Sooss (Austria). D1–D2. Megaclavatula antoniae (Hoernes & Auinger, 1891), Z.PAL.U.W., No BkKG928, Korytnica (Poland). E1–E2. Megaclavatula antoniae (Hoernes & Auinger, 1891), NHMW 1949/0005/0045, Guntersdorf (Austria). F1–F3. Megaclavatula antoniae (Hoernes & Auinger, 1891), holotype, NHMW 1849/0005/0048, Guntersdorf (Austria). G1–G3. Megaclavatula antoniae (Hoernes & Auinger, 1891), NHMW 1949/0005/0045, Guntersdorf (Austria).