Granulolabium plicatum (Bruguière, 1792), Lozouet et al., 2001

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 : 131-134

Granulolabium plicatum (Bruguière, 1792)

Figs 26A–J, 25F

* Cerithium plicatum — Bruguière 1792: 488

Cerithium plicatum Bruguière — Lamarck 1804: 345, pl. 14, fig. 2.

Cerithium plicatum Bruguière — Brongniart 1823: 71, pl. 6, fig. 12.

Cerithium plicatum Bruguière var B. Grateloup— Grateloup 1846: pl. 18, fig. 19.

C [erithium]. plicatum Dub. — Hauer 1837: 419.

[ Cerithium ] plicatum Brug. — Ĥrnes 1848: 21.

Cerithium plicatum Brug. — Ĥrnes 1855: 400, pl. 42, figs 6a–b.

Cerithium Sandbergeri Guemb. —Ģmbel 1861: 754.

[ Cerithium ] elegans Desh. — Fuchs 1868: 217 [non Deshayes, 1824].

[ Cerithium ] plicatum Brug. var. enodosum Sandb. — Fuchs 1868: 217 [? non Sandberger, 1859]

[ Cerithium ] plicatum Brug. var. intermedium Sandb. — Fuchs 1868: 217 [? non Sandberger, 1859].

[ Cerithium ] plicatum Brug. var. multinodosum Sandb. — Fuchs 1868: 217 [? non Sandberger, 1859].

[ Cerithium ] plicatum Brug. var. pustulatum Sandb. — Fuchs 1868: 217 [? non Sandberger, 1859].

[ Potamides (Granulolabium) plicatum (Brug.) ] var. vindobonensis Sacc. — Sacco 1895: 58.

[ Potamides ] G [ranulolabium]. plicatum var. italoturrita Sacc. — Sacco 1895: 58, pl. 3, fig. 45.

[ Potamides ] G [ranulolabium]. plicatum var. perlateplicata Sacc. — Sacco 1895: 58, pl. 3, fig. 46.

Potamides papillatus Sandberger — Penecke 1897: 54, pl. 3, figs 1–3.

Potamides plicatus Brug. — Wolff 1897: 270, pl. 25, fig. 29.

Cerithium bavaricum v. Ģmbel—Ģmbel in Wolff 1897: 269, pl. 26, fig. 6 [nov. nom. pro Cerithium sandbergeri Ģmbel, 1861].

Potamides plicatus Brug. —B̂ckh 1899: 32, pl. 9, fig. 13.

Cerithium (Granulolabium) plicatum Brug. var. papillata Sandb. — Schaffer 1912: 151, pl. 51, figs 36–40 [? non Sandberger, 1859].

Cerithium (Granulolabium) plicatum Brug. var. trinodosa Schff. — Schaffer 1912: 152, pl. 51, figs 36–40 [? non Deshayes 1866].

Cerithium (Granulolabium) plicatum Brug. var. Moldensis Schff.— Schaffer 1912: 153, pl. 51, figs 46–47.

Cerithium (Granulolabium) inaequinodosum Schff. — Schaffer 1912: 153, pl. 51, figs 51–53.

Cerithium (Granulolabium) plicatum Brug. var. quinquenodosa Schff. — Schaffer 1912: 153, pl. 51, figs 44–45.

Cerithium (Clava) bidentatum Defr. var. abbreviata Schff. — Schaffer 1912: 156, pl. 52, fig. 7 [non Leckenby, 1858].

Cerithium (Granulolabium) plicatum Brug. var. papillata Sandb. — Schaffer 1913: 110, pl. 5, fig. 8 [? non Sandberger, 1859].

Cerithium (Granulolabium) plicatum Brug. var. trinodosa Schff. — Schaffer 1913: 111, pl. 5, fig. 9 [non Deshayes 1866].

Cerithium bavaricum Ģmb.— Weithofer 1917: 34.

Cerithium plicatum Bruguière — Favre 1918: pl. 9, figs 164a–b.

Pirenella plicata (Bruguière) — Cossmann & Peyrot 1922: 267, pl. 5, figs 99–101, pl. 6, figs 42–44.

Pirenella inconstans (Basterot) — Cossmann & Peyrot 1922: 270, pl. 6, figs 37–39.

Potamides (Pirenella) plicatus Brug. Gaáli n. sp. —Şmeghy 1922: 33, text-fig. 2.

Clava bidentata schafferi n.n. — Finlay 1927: 495 [nov. nom pro. Cerithium abbreviata Schaffer, 1912 , non Leckenby, 1858].

Cerithium plicatum (Bruguière) — Alimen 1936: 34, 37, etc., pl. 7, figs 10–21.

Pirenella moravica Ĥrnes — Sieber 1937: 483, pl. 24, figs B 1, 2, pl. 25, fig. E1 [= Granulolabium plicatum moravicum ( Ĥrnes, 1855].

Cerithium (Granulolabium) plicatum Brug. var. papillata Sandb. — Munda 1939: 112, pl. 1, fig. 18 [? non Sandberger, 1859].

Pirenella plicata (Bruguière) — Stchépinsky 1946: 138, pl. 30, figs 13–14.

Pirenella plicata (Bruguière) — Anić 1952: 41, pl. 11, fig. 2.

Cerithium plicatum Brug. var. Galeottii Nyst — Kharatishvili 1952: 64 [? non Nyst, 1845].

Cerithium plicatum Brug. — Kharatishvili 1952: 35.

Potamides (Pirenella) plicatus Brug. — Korobkov 1955: plate captions, pl. 28, figs 9–10.

Potamides plicatus Brug. — Strausz 1956: 274, pl. 40, figs 1–15.

P [otamides]. plicata papillata (Sdbgr.)— Sieber 1958: 135.

P [otamides]. plicata moldensis (Schff.)— Sieber 1958: 135.

P [otamides]. inaequinodosa (Schff.)— Sieber 1958: 135.

P [otamides]. plicata quinquenodosa (Schff.) — Sieber 1958: 135.

P [otamides]. plicata trinodosa (Schff.) — Sieber 1958: 135 [non Deshayes 1866].

T [erebralia]. bidentata abbreviata (Schff.) — Sieber 1958: 136 [non Leckenby, 1858].

Pirenella plicata cf. plicata (Brug.) — Seneš 1958: 137.

Pirenella plicata (Bruguière 1889) —Ĥlzl 1958: 191.

Potamides plicatus (Brongn.) — Vadász 1960: 348, pl. 41, fig. 9.

Potamides ( Pyrenella [sic]) plicatus Brug. var. gaáli Şmeghy 1923— Boda 1964: 132.

Cerithium (Pirenella) plicatum Вrug. var. galeottii Nyst — Gamkrelidze et al. 1964: 207 [? non Nyst, 1845].

Pirenella plicata plicata (Bruguière) —Ĥlzl 1965: 261, pl. 3, fig. 10.

Potamides (Pirenella) plicata papillata (Sandberger, 1863) — Ondrejíčková & Seneš 1965: 175, pl. 4, figs 34–39 [? non Sandberger, 1859].

Potamides lamarcki [sic] (Brongn.)— Ondrejíčková & Seneš 1965: 174, pl. 4, figs 40–41 [non Brongniart, 1810].

Potamides (Pirenella) plicatus Bruguière, 1792 — Strausz 1966: 143, pl. 78, fig. 13.

Cerithium plicatum Brug. Galeotti Nyst — Ali-Zade 1968: 73, pl. 1, figs 15a–b [? non Nyst, 1845].

Pirenella plicata papillata (Sandberger, 1863) — Steininger et al. 1971: 377, pl. 3, fig. 13, pl. 4, figs 8–15 [? non Sandberger, 1859].

Pirenella plicata trinodosa Schaffer, 1912 — Steininger et al. 1971: 377, pl. 4, figs 1–2 [non Deshayes, 1866].

Pirenella plicata galeotti (Nyst, 1843) — Steininger et al. 1971: 378, pl. 4, figs 4–7 [? non Nyst, 1845].

Pirenella plicata plicata (Bruguière, 1792) — Moisescu 1972: 89, pl. 29, figs 20a–b.

Pirenella plicata (Bruguière, 1792) — Báldi 1973: 259, pl. 29, fig. 3.

Pirenella plicata (Bruguière, 1792) — Huckriede & Janssen 1973: 192, pl. 1, figs 1, 3–9.

Pirenella plicata bavarica (Ģmbel, 1861)— Janssen 1984: 126.

Pirenella laevissima ( Goldfuss, 1844) — Janssen 1984: 126.

Pirenella plicata (Bruguière) — Gitton et al. 1986: 8, pl. 2, figs 10–11.

Granulolabium plicatum (Bruguière, 1792) — Lozouet 1986: 171, figs 2a–d.

Pirenella plicata (Bruguière, 1792) — Barthelt 1989: 39, pl. 12, fig. 2.

Pirenella binodosa n. sp. — Moisescu 1992a: 13, pl. 1, figs 1–8, text-fig. 1.

Pirenella plicata (Bruguière, 1792) — Moisescu 1992a: 16, pl. 1, figs 9–14, text-fig. 4.

Pirenella trinodosa ( Schaffer, 1912) — Moisescu 1992a: 14, pl. 1, text-figs 2–3 [non Deshayes, 1866].

Pirenella quinquenodosa ( Schaffer, 1912) — Moisescu 1992a: 16, pl. 1, figs 32–34, 37, text-fig. 5.

Pirenella sexanodosa n. sp. — Moisescu 1992a: 16, pl. 1, figs 35–36, 38–40, text-fig. 6.

Pirenella trinodosa ( Schaffer, 1912) — Moisescu 1992b: 19, pl. 3, figs 12a– b [non Deshayes, 1866].

Pirenella plicata papillata (Sandberger) — Popa & Chira 1999: 400, pl. 1, figs 7–9.

Pirenella plicata quinquenodosa Schaffer — Popa & Chira 1999: 400, pl. 1, figs 10–11.

Granulolabium plicatum (Bruguière, 1792) — Lozouet et al. 2001a: 27, pl. 8, figs 1–3.

Granulolabium plicatum (Bruguière, 1792) — Lozouet et al. 2001b: 49, pl. 1, figs 1–2.

Granulolabium plicatum (Bruguière, 1792) — Harzhauser & Kowalke 2001: 362, figs 4.2–5.

Granulolabium plicatum (Bruguière, 1792) — Harzhauser & Mandic 2001: 694, pl. 1, fig. 7.

Pirenella plicata (Bruguière, 1792) — Mikuž 2003: 98, pl. 4, figs 3–4.

Pirenella trinodosa Schaffer, 1912 — Mikuž 2003: 100, pl. 4, fig. 5 [non Deshayes, 1866].

Granulolabium plicatum (Bruguière, 1792) — Harzhauser 2004: 120, pl. 5, figs 1–4.

Granulolabium cf. inaequinodosum ( Schaffer 1912) —Pfister & Wegm̧ller 2007: 91, pl. 7, fig. 7.

Granulolabium plicatum (Bruguière, 1792) — Ýslamoðlu 2008: 266, fig. 6P.

Granulolabium plicatum (Bruguière, 1792) — Esu & Girotti 2010: 159, pl. 5, figs 10–13.

Granulolabium plicatum (Bruguière, 1792) — Moths et al. 2010: 32, pl. 10, fig. 6, pl. 34, fig. 8.

Granulolabium plicatum (Bruguière, 1792) — Lozouet & Maestrati 2012: 278, figs 180/4–6.

Granulolabium plicatum (Bruguière, 1792) — Cluzaud et al. 2014: 226, fig. 171I–K.

Granulolabium plicatum (Bruguière) — Kovács & Vicián 2016: 246, pl. 1, fig. 5.

Granulolabium plicatum (Bruguière, 1792) s. lat. — Marquet et al. 2016: 10, pl. 1, fig. 9.

Granulolabium plicatum (Bruguière, 1792) — Thivaiou et al. 2019: 331, figs 4B1–B3.

non Granulolabium plicatum (Bruguière, 1792) — Vazzana & Cecalupo 2007: 178, pl. 4, fig. m [= Potamides sp. ].

Type material. The syntypes or the holotype were stored in the Museum National d’Histoire Naturelle, Paris (MNHN) but cannot be traced currently (pers. comm. Jean-Michel Pacaud, 11. 12. 2021).

Type locality. La Caunette near Montpellier ( France) .

Stratigraphy. Early Miocene, Aquitanian.

Illustrated material. MNHN.F.A42201, SL: 24.7 mm, MD: 11.0 mm, Auvers-Saint-Georges, Les Sablons ( France), Rupelian, photo: Jacques Mouchart (e-recolnat Project, MNHN), Figs 26A 1 –A 2. MNHN.F.R06865, SL: 35.8 mm, MD: 8.8 mm, Saucats ( France), Aquitanian, photo: Jacques Mouchart (e-recolnat Project, MNHN), Fig

26B. NHMW 2022/0024/0010, SL: 53.1 mm, MD: 13.8 mm, Máriahalom ( Hungary), Figs 26C 1 –C 2. NHMW 1846/0037/0371, SL: 35.8 mm, MD: 9.5 mm, M̂rtersdorf ( Austria), Eggenburgian (Early Miocene), illustrated in Ĥrnes (1855: pl. 42, figs 6a–b), holotype of Granulolabium plicatum vindobonensis Sacco, 1895 , Figs 26D 1 –D 2. NHMW 1912/0001/0015, SL: 34.9 mm, MD: 11.0 mm, Nonndorf ( Austria), Early Miocene, Eggenburgian, lectotype of Cerithium (Granulolabium) plicatum quinquenodosa Schaffer, 1912 , illustrated in Schaffer (1912: pl. 51, fig. 45), Figs 26E 1 –E 2. F /1150, SL: 21.8 mm, MD: 7.9 mm, Krahuletz Museum (Eggenburg, Austria), lectotype of Cerithium (Granulolabium) inaequinodosum Schaffer, 1912 , specimen illustrated in Schaffer (1912: pl. 51, fig. 53), Figs 26F 1 –F 2. F /1149, SL: 25.4 mm, MD: 8.2 mm, paralectotype of Cerithium (Granulolabium) inaequinodosum Schaffer, 1912 , Krahuletz Museum (Eggenburg, Austria), specimen illustrated in Schaffer (1912: pl. 51, fig. 52), Figs 26G 1 –G 2. NHMW 2022/0061/0001, SL: 7.3 mm, MD: 2.8 mm, Mérignac ( France), Early Miocene, Burdigalian, Fig. 26H. NHMW 2022/0062/0001, SL: 18.7 mm, MD: 6.3 mm, Mold ( Austria), Early Miocene, Eggenburgian, Fig. 26I. NHMW 2022/0061/0002, dwarf form, SL: 14.9 mm, MD: 4.6 mm, Mérignac ( France), Early Miocene, Burdigalian Fig. 26J. NHMW 2022/0061/0003, dwarf form, SL: 13.6 mm, MD: 5.9 mm, Mérignac ( France), Early Miocene, Burdigalian, Figs 25F 1 –F 2.

Additional material. Rupelian (early Oligocene): 6 spec., NHMW 1866/0001/0140, Castelgomberto ( Italy) . Egerian (Chattian/Aquitanian): 35. spec., NHMW 2022/0024/0011, Máriahalom ( Hungary). Eggenburgian (Early Miocene): 13 spec., NHMW 1849/0013/0011, Maria Dreieichen ( Austria) ; 18 spec., NHMW 1860/0050/0238, Maria Dreieichen ( Austria) ; 10 spec., NHMW 1934/0001/0079, Maria Dreieichen ( Austria) ; 8 spec., NHMW 1860/0050/0248, Mold ( Austria) ; 5spec.,NHMW1860/0050/0247, Mold ( Austria) ; 5spec.,NHMW1860/0001/0000, Harmersdorf ( Austria) ; 10 spec., NHMW 1866/0040/0034, Nonndorf ( Austria) ; 7 spec., NHMW 1849/0013/0012, Nonndorf ( Austria) ; 1 spec., NHMW 1912/0001/0016, Nonndorf ( Austria) ; 23 spec., NHMW A631, M̂rtersdorf ( Austria) ; 16 spec., NHMW 1860/0050/0359, Kattau ( Austria) ; 1 spec. NHMW 1860/0050/0163, Burgschleinitz ( Austria) ; 2 spec., NHMW 1860/0050/0306, Eggenburg ( Austria) ; 1 spec., NHMW 1860/0050/0280, Ķhnring ( Austria) ; 3 spec., A 637, Bahna ( Romania) ; 8 spec., A 638, Bahna ( Romania) ; 1 spec., NHMW 1860/0007/0004, Zimbor (= Magyarzsombor) ( Romania) . Burdigalian (Early Miocene): 78 spec., NHMW 1856/0029/0078, Bordeaux ( France) ; 11 spec., NHMW 1861/0043/0019, Montpellier ( France) .

Revised description. Shell medium sized, slender conical, of up to 15 teleoconch whorls; apical angle 23–30°. Paucispiral protoconch of 1.5 convex whorls. First teleoconch whorl with two prominent, beaded spiral cords mid-whorl and close to abapical suture. On second whorl, third beaded cord appears above abapical suture, resulting in tripartite sculpture. Abapically, delicate secondary spirals intercalated between primaries and between primaries and adapical and abapical sutures. Later teleoconch whorls with four or five, usually densely spaced, spiral cords of rounded beads, arranged vertically to form more or less pronounced axial ribs. Abapical cords often slightly weaker, with coalescent beads, and frequently partly covered by subsequent whorl. Whorl profile subcylindrical to weakly convex with narrowly incised suture; slightly gradate shells may appear as well. Sculpture highly variable due to intercalations of beaded secondary spiral cords and by bifid primary cords. Last whorl with convex periphery, moderately constricted, with weakly convex base. Base with prominent, weakly beaded or rarely smooth, spiral cords. No varices. Aperture ovoid, moderately wide; outer lip not thickened, opisthocyrt in lateral view; with four to five spiral rows of granules on inner aspect in many specimens; anal canal distinctly incised, often delimited medially by weak parietal swelling; siphonal canal short, moderately incised, wide, only slightly deflected to the left. Columella strongly excavated, smooth. Columellar callus forming broad rim, sharply delimited from base.

Synonyms. Many names erected by Sandberger (1859) for specimens from the Oligocene and Early Miocene of the Mainz Basin have been applied to Paratethyan occurrences. Although we assume that these nominal species in the Mainz Basin might be subjective junior synonyms of Granulolabium plicatum (Bruguière, 1792) , we note that Kadolsky (1995) treated several of these taxa as distinct species, endemic to the Mainz Basin. To avoid incorrect synonymies, we exclude the Mainz Basin records herein from G. plicatum and consequently list Paratethyan record referring to Sandberger (1859) as “ auctores non Sandberger, 1859”.

Potamides (Granulolabium) plicatum vindobonensis was established by Sacco (1895) for the specimen illustrated by Ĥrnes (1855: pl. 42, figs 6a–b). The specimen shows the characteristic spiral rows of granules inside the aperture and a regular sculpture of axially arranged, rather uniform beads. Separation from G. plicatum is not justified. Cerithium (Clava) bidentatum Defr. var. abbreviata was based on a Granulolabium plicatum specimen, which was mistaken for a “ Clava ” (= Terebralia ) by Schaffer (1912). Finlay (1927: 495) recognized that Cerithium abbreviata Schaffer, 1912 was preoccupied by Leckenby (1858: 13) and proposed Clava schafferi as replacement name, which thus is also a secondary junior synonym of G. plicatum . Cerithium (Granulolabium) inaequinodosum Schaffer, 1912 is a morph with prominent secondary spiral cords on early teleoconch whorls. Later teleoconch whorls develop three prominent, beaded adapical spiral cords and several weaker, nearly smooth spiral cords below. The specimens co-occur with typical specimens of G. plicatum , and we consider this species as subjective junior synonym of G. plicatum . Pirenella binodosa Moisescu 1992a is a stout morph of G. plicatum with two spiral rows of beads on the spire whorls. Pirenella sexanodosa Moisescu 1992a is characterized by a weak sixth spiral cord. Cerithium (Granulolabium) trinodosa Schaffer, 1912 is a morph with three dominant spiral cords. Potamides (Pirenella) plicatus gaali Şmeghy, 1922 is another morph with prominently beaded adapical spiral cords and finely granulated abapical spiral cords. Cerithium sandbergeri Ģmbel, 1861 is based on a small specimen with predominant spiral sculpture on late teleoconch whorls. Wolff (1897: 269) emphasized that only few specimens were available of which only one was well preserved. Therefore, we tentatively consider it a rare morph of G. plicatum with reduced beads.

Discussion. The Eggenburgian occurrences display moderate variability and agree well with specimens from the Early Miocene of the Aquitaine Basin. Therefore, both occurrences are treated as conspecific herein.

Distribution. This species is very widespread during the Oligocene and Early Miocene, when it is documented from the North Sea, the northeastern Atlantic, the Paratethys Sea and the Proto-Mediterranean Sea. During the Rupelian, it seems to have even entered the early Indian Ocean and was described by Harzhauser (2007) from the Sultanate of Oman. Typical representatives of this species disappeared from the Paratethys after the Eggenburgian (early Burdigalian). During the Karpatian (late Burdigalian) it was succeeded by Granulolabium moravicum ( Ĥrnes, 1855) .

Occurrences in the Oligocene and Early Miocene of the Mainz Basin are distinguished by Kadolsky (1995) as separate species.

Central Paratethys. Egerian (late Oligocene/Early Miocene): North Alpine Foreland Basin: Kaltenbachgraben ( Germany) (Ĥlzl 1965), Kendl, Loosdorf, Melk, Pielach, Zelking ( Austria) ( Harzhauser & Mandic 2001); South Slovakian Basin: Štúrovo ( Slovakia) ( Ondrejíčková & Seneš 1965); Kováčov Basin: Kováčov ( Slovakia) ( Seneš 1958); Esztergom Basin: Esztergom-Szentgyörgymező ( Hungary) ( Kovács & Vicián 2016); Hungarian Paleogene Basin: Nagyegyháza, Csordakút, Mány, Gyermely, Zsámbék, Tök, Szomor, Máriahalom, Felsőörspuszta, Vasztély, Csolnok, Tarján, Keszt̂lc, Solymár, Budafok, Szentendre, Pomáz, Leányfalu, Dunabogdány, Ĝd, D̂m̂s, Diósjenő, Nagyoroszi, Borsosberény, Tolmács, Becske, Eger, Novaj, Mucsony, Sajókaza ( Hungary) ( Báldi 1973); Central Slovenia: Soteska ( Slovenia) ( Mikuž 2003); Transylvanian Basin: Zimbor ( Romania) ( Moisescu 1972); Getic Basin: Muereasca ( Romania) ( Moisescu 1992b). Eggenburgian (Early Miocene): North Alpine Foreland Basin: Kaltenbachgraben, Thalberggraben ( Germany) (Ĥlzl 1958); Burgschleinitz, Eggenburg, Gauderndorf, Harmersdorf, Ķhnring, Maigen, Maria Dreieichen, M̂rtersdorf, Molt, Nonndorf ( Austria) ( Steininger et al. 1971); Central Slovakia: Veľká Čausa ( Slovakia) ( Steininger et al. 1973); Borod Basin: Valea Cetea ( Romania) ( Moisescu 1992a).

Eastern Paratethys. Rupelian (early Oligocene): Transcaucasia: Karatubani ( Kacharava, 1944) ;? Sakaraulian (Early Miocene): Kaspi ( Georgia) ( Kharatishvili 1952) (will need verification).

North Sea Basin. Rupelian to Burdigalian (Oligocene/Early Miocene): North Sea Basin: ( Germany, Belgium) ( Huckriede & Janssen 1973; Moths et al. 2010; Marquet et al. 2016); Oligocene:

Paris Basin. France ( Lozouet & Maestrati 2012).

Hampshire Basin. England ( Munt & Barker 1996).

Northeastern Atlantic. Rupelian to Burdigalian (Oligocene/Early Miocene): Aquitaine Basin ( France) ( Cossmann & Peyrot 1922, Lozouet 1986; Lozouet et al. 2001).

Proto-Mediterranean Sea. Rupelian (early Oligocene): Colli Torinesi ( Italy) ( Sacco 1895); Denizli Basin ( Turkey) ( Ýslamoðlu 2008), Esfahan Basin ( Iran) ( Harzhauser 2004). Chattian–Aquitanian (late Oligocene/Early Miocene): Mesohellenic Basin ( Greece) ( Harzhauser 2004; Thivaiou et al. 2019). Aquitanian–Burdigalian (Early Miocene): Sivas Basin ( Stchepinsky 1946). Rupelian: Duqum ( Sultanate of Oman) (Harzhauser 2007). Aquitanian: southern Italy ( Italy) ( Esu & Girotti 2010).