taxonID	type	description	language	source
03FEF726FFFE4E040FADF94469B997F0.taxon	type_taxon	Type species. Chiton cajetanus Poli, 1791, by subsequent designation (Herrmannsen 1846). Non Lepidopleurus (Carpenter MS) Dall, 1879 (= Lepidozona Pilsbry, 1892).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFE4E040FADF94469B997F0.taxon	distribution	Distribution. Lepidopleurus is known from the Eocene to the Recent, with only a living species known (L. cajetanus), from the Mediterranean Sea and the Atlantic Ocean (from Spain and Portugal south to Morocco and Canary Islands). Lepidopleurus sp. has been cited from the Permian of Malaysia (Gobbett & Hutchison 1973: pl. 10, fig. 9), but we consider this single record doubtful. The genus is recorded with confidence in the Eocene of Ukraine (Bielokrys 2000), Germany and France (Dell’Angelo et al. 2011), upper Eocene / lower Oligocene of U. S. A. (Dell’Angelo et al. 2011), Oligocene of Germany (Janssen 1978), Miocene of Japan (Itoigawa et al. 1977), France, Italy and Paratethys (Dell’Angelo et al. 2007 a, 2015 a, 2016, 2018 a; Studencka & Dulai 2010), Pliocene of Spain (Almera 1894; Dell’Angelo et al. 2004), France (Dell’Angelo et al. 2018 b) and Italy (Dell’Angelo et al. 2001 a, 2013)], Pleistocene of Italy (Sacco 1897; Sabelli & Taviani 1979; Chirli 2004), Greece (Garilli et al. 2005; Koskeridou et al. 2009), and Tunisia (Castany et al. 1956).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFE4E040FADF94469B997F0.taxon	discussion	Remarks. The relationships between Lepidopleurus Risso, 1826 and Leptochiton Gray, 1847, have been long debated (Pilsbry 1892; Ferreira 1979; Dell’Angelo & Palazzi 1989, 1991), and even the molecular phylogenetic study by Sigwart et al. (2011) has not solved all problems. We consider Lepidopleurus (“ valves solid, tegmentum heavily sculptured of heavy concentric ridges ”) and Leptochiton (“ valves thin, tegmentum finely granulose, granules of equal size ”) to be distinct genera (Kaas & Van Belle 1985 a). The main morphological characteristics of the Lepidopleurus spp. considered in the present study are reported in Tab. 1.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFB4E050FADFC8C6868912C.taxon	description	Fig. 1	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFB4E050FADFC8C6868912C.taxon	materials_examined	Type material. Holotype: MHNBx 2017.7.1, intermediate valve, width 12 mm (Figs 1 A – B). Paratype: MHNBx 2017.7.2, tail valve, width 9.4 mm (Figs 1 E – F). Type locality. Mineur (France). Type stage. Oligocene, Chattian. Material examined. Upper Oligocene (Chattian): France: Abesse: 8 valves (AC, DA, Figs 1 C-D, 1 G-H), Mineur: type material plus 12 valves (AC). Maximum width of the valves: -- / 14 / 9.4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFB4E050FADFC8C6868912C.taxon	description	Description. Valves solid. Head valve unknown. Intermediate valves broadly rectangular (W / L = 2.42), wide, rounded in anterior profile, moderately elevated (H / W = 0.31 – 0.47), anterior and posterior margins straight, side margins rounded, apex inconspicuous, lateral areas raised. Tail valve semicircular, elevated, width ca. two times the length (W / L = 2.04), anterior margin straight, mucro flat, in anterior position, antemucronal slope slightly convex, postmucronal slope slightly concave just behind mucro. Tegmentum rough, space between striae of granules large, no evidence of growth lines. LA and PMA sculptured with radial chains of large and irregular roundish granules (LA 4 – 6, PMA 40), tending to split near side margins. CA and AMA sculptured with longitudinal chains of large and irregular roundish granules (CA ca. 33), tending to branching and anastomosing in posterior margin near apex. Articulamentum without insertion laminae, apophyses triangular in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFB4E050FADFC8C6868912C.taxon	discussion	Remarks. This species was attributed to the genus Leptochiton Gray, 1847 in the original description, but the robustness and dimensions of the valves are more compatible with Lepidopleurus Risso, 1826. The fossil record of Lepidopleurus aturriensis (Dell’Angelo, Lesport, Cluzaud & Sosso, 2018) is limited to the Oligocene (Chattian) of France (Dell’Angelo et al. 2018 a). The shape and sculpture of the valves are well characterized and display little variability. A single intermediate valve from Mineur is flatter, less rounded, almost subcarinated, H / W = 0.31 (Dell’Angelo et al. 2018 a: fig. 3 S – T). Comparisons. Lepidopleurus aturriensis presents a coarse sculpture, at first glance quite similar to some Lepidopleurus spp. from the Oligocene of Germany, e. g., L. benoisti (de Rochebrune, 1882) and L. vigifer (Sandberger, 1859). Lepidopleurus aturriensis differs from L. benoisti by the higher intermediate valves (H / W = 0.31 – 0.47 vs. 0.33 – 0.36), and the more regular and less coarse sculpture, with no evidence of terraced ribs in LA and PMA; Lepidopleurus virgifer differ from L. aturriensis by the very irregular and coarse sculpture, particularly in the lateral areas of intermediate valves and postmucronal area of tail valve, sculptured with numerous diverging, coarsely granulated radial ribs, crossed by coarse growth ridges.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFB4E050FADFC8C6868912C.taxon	distribution	Distribution. Upper Oligocene: northeastern Atlantic (Chattian): Aquitaine Basin, France: Abesse, Mineur (Dell’Angelo et al. 2018 a, 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFA4E070FADFBF06F049140.taxon	description	Fig. 2	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFA4E070FADFBF06F049140.taxon	description	Dell’Angelo & Palazzi 1989). Chiton benoisti; Cossmann & Peyrot 1919, p. 33, pl. 2, figs 23 – 27; Forli & Guerrini 2022, fig. 11.18 (6). Lepidopleurus subcajetanus; Laghi 1977, p. 99, pl. 1, fig. 21. Lepidopleurus benoisti; Dell’Angelo et al. 2015 a, p. 222, pl. 2, figs 1 – 13, 18 – 20; Dell’Angelo et al. 2018 a, p. 12, figs 2 J – U, 3 A – F; Dell’Angelo et al. 2020 b, p. 52, tab. 9; Stein et al. 2021, p. 55, textfig. 6, pl. 6, fig. 4.? Lepidopleurus sp.; Moths et al. 2010, p. 30, pl. 4, fig. 3; Stein et al. 2016, p. 176 (fide Stein et al. 2021).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFA4E070FADFBF06F049140.taxon	materials_examined	Type material. Syntypes MHNBx 2009. TY. P. 306.0, two tail valves from the Benoist’s collection (width respectively 13.63 and 7.04 mm). Type locality. Mérignac (Gironde, France). Type stage. Lower Miocene, Aquitanian / Burdigalian. Material examined. Upper Oligocene: France, Aquitaine Basin (Chattian): Abesse: 15 valves (AC, JVC, Fig. 2 A – D). Lower Miocene: France, Aquitaine Basin (Aquitanian): Maïnot: 1 valve (PR); Aquitaine Basin (Burdigalian): Cabanes: 1 valve (JFL), Carrière Vives: 5 valves (AC, BD 261, JFL, PR), Gamachot: 3 valves (AC, JFL), la Flotte: 1 valve (AC), Lahitet: 1 valve (AC), Lorient: 7 valves (AC, JFL, PR), Peloua: 33 valves (AC, JFL, JVC, PR, Fig. 2 E – F), Petit Bargues: 1 valve (PR), Pont St Martin: 23 valves (JFL, PR, Fig. 2 G – H). Italy: Torino Hills (Burdigalian): Valle Ceppi: 5 valves (PMRSN BS. 105.02.001 – 003, MZB 32008 – 32009). Middle Miocene: France, Aquitaine Basin (Langhian): Carrière Gélis: 1 valve (JFL); Aquitaine Basin (Serravallian): Orthez: 1 valve (BD 262). Maximum width of the valves: 15.7 / 18 / 18 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFA4E070FADFBF06F049140.taxon	description	Description. Valves solid. Head valve large, semicircular, posterior margin widely V-shaped, slope almost straight or slightly convex, interrupted by profile of concentric, terraced ribs. Intermediate valves wide, broadly rectangular (W / L = 2.45), rounded in anterior profile, moderately elevated (H / W = 0.33 – 0.36), anterior and posterior margins straight, side margins rounded, apex inconspicuous, lateral areas not very raised. Tail valve semicircular (W / L = 1.64 – 1.76), elevate, mucro flat, in anterior position, antemucronal slope almost straight or slightly convex, postmucronal slope almost straight or slightly concave. Tegmentum coarse, space between striae of granules reduced. HV, LA and PMA sculptured with numerous and very irregular radial chains of roundish granules, tending to branching and anastomosing, intersected by many and well evident concentric terraced ribs (up to 5 – 6 in LA). CA and AMA sculptured with many longitudinal chains of roundish granules (ca 100 or more in CA), almost regular in central part, much irregular and tending to branching and anastomosing in lateral parts, near side margins, with concentric ribs inconspicuous, barely visible (and not terraced). Articulamentum without insertion laminae, apophyses narrow, rounded-triangular, widely projected in intermediate valves, larger but not completely preserved in tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFA4E070FADFBF06F049140.taxon	discussion	Remarks. Dell’Angelo et al. (2015 a) examined Chiton subcajetanus d’Orbigny, 1852 sensu Sacco 1897 and the related species (Gymnoplax orbignyi de Rochebrune, 1882 and G. benoisti de Rochebrune, 1882); they consider Chiton subcajetanus a nomen nudum [no diagnosis, nec figuration in Bonelli (1824), Sismonda (1842, 1847) and d’Orbigny (1852)], and Lepidopleurus benoisti (de Rochebrune, 1882) the correct taxon to attribute to this species (including the large valves of L. decoratus of Šulc). Šulc considered a tail valve (width of 18 mm) from the Middle Miocene of Pötzleinsdorf, Austria and the valves illustrated by Sacco (as Middendorffia subcajetana) as a very large form of Lepidopleurus decoratus (Reuss, 1860). In the Šulc collection housed at the NHMW, large tail valves — reported in the “ Material Examined ” section for Lepidopleurus cajetanus (Poli, 1791) — are absent. We concur with Laghi (1977), Bałuk (1984), Studencka & Dulai (2010), and other authors in considering L. decoratus conspecific with L. cajetanus. Accordingly, we believe that records of “ subcajetanus ” or “ decoratus ” from the Paratethys should be attributed to the morphospecies Lepidopleurus cajetanus. Many valves of Lepidopleurus benoisti were described by Dell’Angelo et al. (2018 a) from the Aquitaine Basin of France, in sediments from upper Oligocene to Middle Miocene, and this has allowed the redescription of this species, first known only for few valves from the lower Miocene of France (de Rochebrune 1882; Cossmann & Peyrot 1919) and Italy (Turin hills: Sacco 1897; Dell’Angelo et al. 2015 a). The valves examined from the Aquitaine Basin of France mainly include tail valves, and it is possible to see a certain range of variability. The mucro is more pronounced in juvenile specimens, and the postmucronal slope decidedly concave (Dell’Angelo et al. 2018 a: fig. 3 F, width 4.2 mm), while the position of the mucro remains in anterior position in fully developed specimens, on the contrary of what happens for Lepidopleurus cajetanus. In fact tail valves of Lepidopleurus benoisti are similar to those of L. cajetanus at the second stage of growth (see above), but with larger dimensions, larger than those reached by L. cajetanus at the last stage of growth, the mucro in anterior position, and less evident concentric terraced ribs in the postmucronal area (real concentric folds in L. cajetanus). The report of Chiton miocenicus Michelotti, 1847 by Cossmann & Peyrot (1919: 32, pl. 2, figs 21 – 22) can be attributed to Lepidopleurus benoisti, as already indicated by Dell’Angelo & Palazzi (1989: 55). Cossmann & Peyrot found three tail valves from Salies-de-Béarn (coll. Degrange-Touzin), one of which figured in Cossmann & Peyrot (1919: pl. 2, figs 21 – 22); Dell’Angelo et al. (2018 a) examined these valves preserved at MHNBx and confirmed the attribution to L. benoisti. Comparisons. This species is superficially similar to the recently described Lepidopleurus pseudobenoisti Dell’Angelo et al., 2018 (see below), from which it differs by the different shape of the tail valve with the very pronounced mucro (compare Figs 2 F, 2 H with Figs 5 B, 5 L), and also by the coarser and very variable sculpture and the much more raised lateral areas of intermediate valves in L. pseudobenoisti. These two species cannot be separated based on their head valve (shape or sculpture). Another similar species is Lepidopleurus virgifer (Sandberger, 1859) from the Middle Oligocene (Rupelian) of Waldböckelheim, Germany (see below). The shape and the ornamentation of the valves are similar, but the sculpture is coarser in L. benoisti, with more pronounced growth lines, and the size of the valves is greater.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFFA4E070FADFBF06F049140.taxon	distribution	Distribution. Upper Oligocene: northeastern Atlantic (Chattian): Aquitaine Basin, France: Abesse (Dell’Angelo et al. 2018 a). Upper Oligocene to lower Miocene: North Europe, Germany: Neetze (Stein et al. 2021). Lower Miocene: North Europe, Germany: Werder (Moths et al. 2010; Stein et al. 2016); Northeastern Atlantic (Aquitanian-Burdigalian): Aquitaine Basin, France: Cabanes, Carrière Vives, Gamachot, la Flotte, Lahitet, Lorient, Maïnot, Peloua, Petit Bargues, Pont St Martin (Cossmann & Peyrot 1919; Dell’Angelo et al. 2018 a); Proto-Mediterranean Sea (Burdigalian): N. Italy: Torino Hill (Sacco 1897; Dell’Angelo et al. 2015 a). Middle Miocene: northeastern Atlantic, France, Aquitaine Basin: Carrière Gélis (Langhian), Orthez (Serravallian) (Dell’Angelo et al. 2018 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF84E0B0FADFB5C69CD9420.taxon	description	Fig. 3	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF84E0B0FADFB5C69CD9420.taxon	materials_examined	Type material. Probably lost. Lectotype designated by Dell’Angelo & Palazzi (1989), specimen figured by Poli (1791: pl. 4, fig. 1). Type locality. Gaeta (Italy). Material examined. Lower Miocene: France, Aquitaine Basin (Burdigalian): Pont St Martin: 1 valve (PR). Middle Miocene: France, Northeastern Atlantic (Langhian): Pouyouet: 1 valve (PR); Central Paratethys: Austria: Grund: 1 valve (NHMW 2010 / 0256 / 0020), Niederkreuzstetten: 1 valve (NHMW 2010 / 0256 / 0002), Niederleis: 2 valves (NHMW 1863 / 0015 / 0860), Pötzleinsdorf: 163 valves (NHMW 1859 / 0027 / 0060 – 0061, Figs 3 J – K, 1859 / 0038 / 0191, Figs 3 F – I, 1861 / 0028 / 0079, 1865 / 0001 / 0987), Speising: 1 valve (NHMW 1860 / 0028 / 0108); Romania: Bujtur: 2 valves (NHMW 1863 / 0012 / 0116), Kostej: 5 valves (BD 263, NHMW 1867 / 0019 / 0345), Lapugy: 14 valves (BD 264, NHMW 2010 / 0256 / 0004, 2010 / 0256 / 0024); Hungary: Bánd: 78 valves (BD 265), Letkés: 6 valves (BD 266); Eastern Paratethys: Ukraine: Horodok: 4 valves (BD 267), Varovtsi: 1 valve (BD 268), Velyka Levada: 1 valve (BD 269). Upper Miocene: France, Ligerian Basin (Tortonian): Moulin Pochas: 20 valves (PR); Anjou: Saint-Clément-de-la-Place: 79 valves (MNHN. F. A 67042 – A 67046, NHMW 2017 / 0108 / 0001, RGM. 1008397, BD 129, Figs 3 O – P), Renauleau: 17 valves (MNHN. F. A 67047 – A 67048, NHMW 2017 / 0108 / 0002, BD 130), Sceaux d’Anjou: 3 valves (RGM. 1008442, RGM. 1008447); Italy, Po Basin (Tortonian): Borelli: 38 valves (BD 270, Figs 3 A – B, MGPT PU 135037, MZB 32003, MZB 32007, PG), Montegibbio: 21 valves (BD 271, MZB 32004, MZB 32006), Rio di Bocca d’Asino: 58 valves (MZB 32001 – 32002, BD 272, Figs 3 M – N, PG), Vigoleno: 1 valve (MZB 32044), Villa Monti: 79 valves (BD 273, MZB 32005, PG). Lower Pliocene: Italy: Liguria: Borzoli: 1240 valves (BD 274, MSNG, Figs 3 C – E, MZB 45692, MZB 45695, MZB 45697 – 45698), Bussana: 269 valves (BD 275, MZB 45690 – 45691, PG, SR), Garlenda: 67 valves (MP), Genova Sestri: 11 valves (BD 276), Rio S. Antonino: 2280 valves (BD 277, MP, MZB 45693 – 45694, MZB 45696, PG), Rio Torsero: 2 valves (BD 278, MP), Caranchi: 5 valves (MP), Salea: 1 valve (BD 279), Zinola: 13 valves (BD 280). Pliocene: Spain: Estepona: 7 valves (BD 281); Italy: Piedmont: Vintebbio: 11 valves (BD 282); Tuscany: Castiglioncello del Trinoro: 2 valves (BD 283), Montenero: 11 valves (BD 284), Serre di Rapolano: 60 valves (BD 285), Villa Banfi: 3 valves (BD 286); Emilia-Romagna: Bacedasco: 11 valves (BD 287), Gagliardella: 3 valves (BD 288), Sariano: 1 valve (BD 289), Vernasca: 14 valves (BD 290); Sicily: Trappeto: 11 valves (BD 291). Pleistocene: France: Corsica, Patrimonio: 8 valves (BD 292); Italy: Tuscany: Cisternino: 32 valves (BD 293), Riparbella: 3 valves (BD 294); Emilia-Romagna: Torrente Stirone: 20 valves (BD 295); Tuscany: Fauglia: 2 valves (BD 296); Puglia: Gallipoli: 2 valves (BD 297), Punta Penne: 3 valves (BD 298); Calabria: Archi S. Francesco: 8 valves (BD 299), Gallina: 3 valves (BD 300), Le Castella: 20 valves (BD 301), Musalà: 8 valves (BD 302), Pecoraro: 3 valves (BD 303), Pezzo: 13 valves (BD 304), S. Maria di Catanzaro: 3 valves (BD 305), Stalettì: 3 valves (BD 306), Terreti: 4 valves (BD 307); Sicily: Calderà: 1 valve (BD 308), Capo Milazzo: 8 valves (BD 309), Grammichele: 3 valves (BD 310), Messina: 1 valve (BD 311); Greece: Kyllini: 2 valves (DGUP, Fig. 3 L). Maximum width of the valves: 8.3 / 10 / 8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF84E0B0FADFB5C69CD9420.taxon	description	Description. Valves solid. Head valve semicircular, posterior margin almost straight, front slope weakly concave, interrupted by profile of concentric, terraced ribs. Intermediate valves broadly rectangular, with a very variable width / length ratio (H / W = 2.41 – 3.60), rounded in anterior profile, moderately elevated (H / W = 0.28 – 0.40), anterior margin variable, from concave to straight or convex, lateral margins almost straight or slightly rounded, posterior margin straight with apex not evident, lateral areas triangular, strongly elevated, starting from near the apex to neighbouring the lateral margin, moderately angled (30 – 45 °) to the posterior margin. Tail valve semicircular, shape strongly variable with size, anterior margin straight or little convex, mucro evident and variable with size from subcentral to forwardly produced, antemucronal slope almost straight, postmucronal slope variable with size. Tegmentum coarse, very variable. HV, LA and PMA sculptured with strong, concentric, terraced ribs, spaced from each other. CA and AMA sculptured with longitudinal chains of granules, somewhat branching or anastomosing, transversally intersected by much thinner cords that give a pitted appearance, more evident in younger individuals. Granules of irregular shape, from roundish to polygonal, up to 100 µm long, elevated, united with each other, with one subcentral megalaesthete and up to 10 micraesthetes placed on both sides of granules. Articulamentum without insertion laminae, apophyses small, triangular, divided by a wide jugal sinus.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF84E0B0FADFB5C69CD9420.taxon	discussion	Remarks. The species displays considerable morphological variations from juvenile stages to adulthood. Laghi (1977: fig. 3 a-b) documented that the mucro of tail valves is almost central in juvenile specimens, moving backward (up to the end of the valve) with growth, due to the bending of the posterior area on the ventral side. The first postlarval stage of juvenile Lepidopleurus cajetanus (Poli, 1791) is characterized by a leptochitonid-like sculpture, i. e. with small cords of tubercles but without the presence of “ steps ” (terraced ribs) on the valves and with the mucro anterior or pre-central. Later, the typical ‘ step’ sculpture appears but the mucro is still subcentral and the small cords of tubercles are not cancelled (Figs 3 L). In the final stage, the small cords are almost indistinguishable in the “ step ” zone, while the mucro is situated more posteriorly and the outline is more triangular (Fig. 3 O – P). These features have well described and illustrated by Laghi (1977: fig. 3 a – b), and Dell’Angelo et al. (2013: pl. 1, figs F – G; 2015 a: pl. 1, figs 9 – 12). The morphology of intermediate valves attributed in the literature to Lepidopleurus cajetanus, consists, in fact, of two distinct types of LA. The first type (Lepidopleurus cajetanus), has the starting point of the lateral area with the concentric terraced ribs near the apex, more regular, slightly angulated on the posterior margin (30 – 45 °), rounded in anterior profile but the valve is not very elevated (H / W = 0.28 – 0.40). The second type shows the starting point of the lateral area with the concentric terraced ribs neighbouring the lateral margin (not near the apex), much more angled at the posterior margin, from 50 – 60 ° upwards up to almost perpendicular, rounded in anterior profile but more elevated (H / W = 0.40 – 0.64), in addition to other minor features (see below). These remarkable differences may well account for representing two distinct taxa, which can, however, be discriminated only upon intermediate valves. A new species, Lepidopleurus pseudocajetanus sp. nov. is described below. Another remarkable example of variation in the sculpture of the tegmentum can be evidenced in CA and AMA, normally with longitudinal chains of granules (Fig. 3 C), somewhat branching or anastomosing, transversally intersected by thinner cords that give a pitted appearance (Fig. 3 M). The longitudinal chains may be irregularly directed towards the sides in some cases (Fig. 3 L), but the sculpture of “ parallel ” chains is always well evident. This type of sculpture is prevalent in Recent specimens (see Dell’Angelo & Smriglio, 1999: pl. 6, fig. E, pl. 7, figs K, L), but also in valves from Pleistocene and Pliocene (see Garilli et al. 2005: pl. 1, fig. 1 from the Pleistocene of Kyllini, Greece; Dell’Angelo et al. 2013: pl. 1, fig. B from the Pliocene of Liguria). The sculpture prevailing in Miocene specimens is more irregular and rougher, shaped by groups of granules branching longitudinally many times, and giving an appearance without evidence of longitudinal chains (Fig. 3 J). Furthermore, the intersections by thinner cords are not present. The variability is very large, with multiple branching occurring only in a part of the central area of some valves. Although less frequently, such irregular sculpture may also occur in Pliocene to modern valves. The great variability of the valves and the remarkable variations in morphology with age found in Lepidopleurus cajetanus, is confirmed by the check of the valves of L. decoratus (Reuss, 1860) present in the Šulc collection. The same degree of variability is also present in valves from the Paratethys (Dulai 2005; Studencka & Dulai 2010), as already reported by Reuss and Šulc (see Reuss, pl. 8, fig. 7). We can therefore confirm that L. decoratus must be considered a synonym of L. cajetanus. Comparisons. The characteristics of the tegmentum sculpture (with the presence of strong, concentric, terraced ribs on HV, LA, PMA) make this species easily identifiable and unmistakable.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF84E0B0FADFB5C69CD9420.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Pont St Martin (Dell’Angelo et al. 2018 a). Middle Miocene: northeastern Atlantic (Langhian): Aquitaine Basin, France: Pouyouet (Dell’Angelo et al. 2018 a); Central Paratethys (Langhian-Serravallian): Austria: Grund, Niederkreuzstetten, Niederleis, Pötzleinsdorf, Speising, Steinabrunn (Šulc 1934; this study), Czech Republic: Borač, Kninice, Rudoltice (Šulc 1934), Slovakia: Dubová (Ruman & Hudácková 2015), Poland: Korytnica (Bałuk, 1971, 1984), Romania: Bujtur, Coştei, Lăpugiu de Sus (Zilch 1934; Dell’Angelo et al. 2007 a; this study), Hungary: Bánd, Letkés (Dulai 2005, 2025 b; this study); Eastern Paratethys: Ukraine: Horodok, Podhorce, Szuszkovce, Varovtsy, Velyka Levada (Studencka & Dulai 2010; this study). Upper Miocene: northeastern Atlantic: Ligerian Basin, France: Moulin Pochas, Saint-Clément-de-la-Place, Renauleau, Sceaux d’Anjou (Dell’Angelo et al. 2018 a, 2018 b); Proto-Mediterranean Sea: Po Basin, N Italy: Borelli, Montegibbio, Rio di Bocca d’Asino, Vigoleno, Villa Monti (Laghi 1977; Dell’Angelo et al. 1999, 2015 a). Lower Pliocene: central Mediterranean, Italy: many localities in Liguria (Laghi 1977; Dell’Angelo et al. 2001 a, 2013; Chirli 2004). Pliocene: Western Mediterranean, Spain: Estepona (Dell’Angelo et al. 2004); France: Biot (B. D. D. 1882); Italy: many localities in Piedmont, Tuscany, Emilia-Romagna, Sicily. Upper Pliocene to upper Pleistocene: central Mediterranean, Greece: Rhodes (Koskeridou et al. 2009). Pleistocene: central Mediterranean, Italy: many localities in Tuscany, Puglia, Calabria, Sicily (Sabelli & Taviani 1979; Dell’Angelo & Palazzi 1989; Dell’Angelo et al. 2001 a); Greece: Kyllini (Garilli et al. 2005), Tunisia: Monastir (Castany et al. 1956). Recent: Atlantic Ocean, from Spain and Portugal (Borja 1987; Consolado Macedo et al. 1999; Urgorri et al. 2017) south to Morocco (Pallary 1920) and Canary Islands (Kaas 1991; Hernández & Rolán 2011) and Berlengas Archipelago (Pisani Burnay 1986). Mediterranean Sea: Spain and Balearic Islands (Altaba 1993; Moreno & Gofas 2011); France (B. D. D. 1882; Leloup 1934); Italy: Lampedusa Island (Spada et al. 1973) and many localities (Leloup & Volz 1938; Dell’Angelo & Smriglio 1999); Croatia (Leloup & Volz 1938); Malta (Mifsud et al. 1990); Greece and Aegean Sea Islands (Strack 1988; Koukouras & Karachle 2005); Turkey (Demir 2003; Ozturk et al. 2014); Israel (Barash & Danin 1977); Libia (Monterosato 1923); Algeria: Orano (Pallary 1900).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF44E0B0FADFEFC6BBD920A.taxon	description	Fig. 4	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF44E0B0FADFEFC6BBD920A.taxon	materials_examined	Type material. Holotype: MNHN. F. A 67060, tail valve, width 14 mm (Figs 4 A – D). Paratype: MNHN. F. A 67061, tail valve, width 10 mm. Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Upper Miocene: France: Saint-Clément-de-la-Place: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF44E0B0FADFEFC6BBD920A.taxon	description	Description. Valves solid. Head and intermediate valves unknown. Tail valve semicircular, elevated, width slightly less than twice length (W / L = 1.77), anterior margin slightly convex, mucro pronounced, in anterior position, antemucronal slope convex, postmucronal slope slightly concave just under mucro, interrupted by profile of concentric terraced ribs. Tegmentum fine, more regular. PMA sculptured with radial chains of granules, just over 80 near mucro, tending to branch and multiply towards posterior margin, intersected by numerous concentric ribs. AA sculptured with ca. 110 regular longitudinal chains of roundish granules, united with each other. Articulamentum without insertion laminae, apophyses large.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF44E0B0FADFEFC6BBD920A.taxon	discussion	Remarks. The species is known only from the two tail valves from Saint-Clément-de-la-Place (France) described by Dell’Angelo et al. (2018 b). Comparisons. These valves differ from tail valves of Lepidopleurus pseudobenoisti Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 in the regularity (no trace of branching and anastomosing) and the greater number (110 vs. 60) of the longitudinal chains of granules in AMA, the less concave postmucronal slope, and conspicuously finer sculpture. See Tab. 1 for a comparison with the Lepidopleurus spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF44E0B0FADFEFC6BBD920A.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Anjou, France: Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF34E0D0FADFF05697E9164.taxon	description	Fig. 5	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF34E0D0FADFF05697E9164.taxon	materials_examined	Type material. Holotype MNHN. F. A 67049, tail valve (Figs 5 A – B). Paratypes: MNHN. F. A 67050 – A 67052 (3 valves); NHMW 2017 / 0108 / 0003 – 0005 (3 valves); RGM. 1008393 – 1008395 (3 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Upper Miocene: France: Saint-Clément-de-la-Place: type material plus 697 valves (MNHN. F. A 67053 – A 67059, Figs 5 E – F, 5 I – J), NHMW 2017 / 0108 / 0006, RGM. 1008349, RGM 1008396, RGM 1008414, RGM 1008427, RGM 1008440, BD 131, Figs 5 C – D, G – H, K – L), Sceaux d’Anjou: 2 valves (RGM 1008443, BD 132). Maximum width of the valves: 15.5 / 26.2 / 18.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF34E0D0FADFF05697E9164.taxon	description	Description. Valves solid. Head valve large, semicircular, posterior margin widely V-shaped, slope almost straight or slightly convex interrupted by profile of concentric, terraced ribs. Intermediate valves wide, broadly rectangular, width more than three times the length (W / L = 3.16 – 3.44), rounded in anterior profile, moderately elevated (H / W = 0.25 – 0.33), anterior margin slightly convex, side margins rounded, posterior margin straight, apex inconspicuous, lateral areas strongly raised. Tail valve semicircular, elevated, width slightly less than twice length (W / L = 1.72 – 1.85), anterior margin straight and slightly convex in jugal area, mucro strongly pronounced, in anterior position, antemucronal slope decidedly convex, postmucronal slope concave just under mucro, interrupted by profile of concentric, terraced ribs. Tegmentum coarse, space between striae of granules large. HV, LA and PMA sculptured with numerous and strongly irregular branching or anastomosing radial chains of granules, intersected by some prominent concentric, terraced ribs (up to 5 – 6 in LA). CA and AMA sculptured with numerous longitudinal chains (ca 60 in CA) of roundish, large granules (diameter up to 100 µm), more regular in central part, markedly irregular and tending to branching and anastomosing laterally near side margins, sculpture on CA even more irregular and coarser than on AMA. Each granule with a central megalaesthete and up to 20 micraesthetes irregularly disposed along margin. Articulamentum without insertion laminae, apophyses narrow, triangular, widely projected on intermediate valves, larger and rounded on tail valves, muscle scars particularly well preserved on tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF34E0D0FADFF05697E9164.taxon	discussion	Remarks. The valves examined are the largest found in the genus Lepidopleurus. In spite of the relative abundance of fossil material, only a few valves preserve their features complete, many missing the apophyses. The sculpture is variable, especially with respect to the longitudinal chains of granules in CA, which tend to branch and anastomose irregularly. It is difficult to count the number of longitudinal chains of granules, ca. 60 counted on the anterior margin. The longitudinal chains in the holotype are more regular, almost parallel and with only a few chains branching and anastomosing in the jugal area (Fig. 5 A). The intermediate valves have irregular sculpture (Dell’Angelo et al. 2018 b). Some valves (Fig. 5 I) display a sculpture in CA in which some longitudinal chains are missing, and a very irregular sculpture in LA. Numerous small valves are interpreted as juvenile specimens of L. pseudobenoisti (Dell’Angelo et al. 2018 b: fig. 2 P – R), showing smaller roundish granules (diameter ca. 65 – 75 µm), with a central megalaesthete and fewer micraesthetes (7 – 10) regularly disposed along the margin. Comparisons. Lepidopleurus pseudobenoisti Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is superficially similar to L. benoisti (de Rochebrune, 1882), but L. pseudobenoisti differs most importantly by the different shape of the tail valve with the strongly pronounced mucro and also by the coarser and more variable sculpture and the considerably more elevated lateral areas of intermediate valves. Another similar species is Lepidopleurus virgifer (Sandberger, 1859) (see below), from which L. pseudobenoisti differs by its smaller size and the shape of the tail valve with a strongly pronounced mucro.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF34E0D0FADFF05697E9164.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Anjou, France: Saint-Clément-de-la-Place, Renauleau, Sceaux d’Anjou (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	description	Fig. 6	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	materials_examined	Type material. Holotype: MSNG 62623, intermediate valve, width 9.5 mm (Figs 6 F – H). Paratype 1: MSNG 62624, intermediate valve, width 8 mm (Figs 6 A – C), from Saint-Clément-de-la-Place, France, Miocene (Tortonian). Paratype 2: MNHN. F. A 98468, intermediate valve, width 6.5 mm (Figs 6 D – E), from Rio S. Antonino (Italy), Pliocene (Zanclean). Paratype 3: SMF 380817, intermediate valve, width 7 mm, from Bussana (Italy), Pliocene (Zanclean). Paratype 4: NHMW 1861 / 0028 / 0079 (as Lepidopleurus decoratus), intermediate valve, width 9 mm (Figs 6 I – L), from Pötzleinsdorf (Austria), Middle Miocene. Type locality. Pezzo, near Villa S. Giovanni (Calabria, Italy). Type stage. Upper Pleistocene, but presumably originated from contiguous deposits of the Lower Pleistocene of bathyal facies.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	etymology	Etymology. The name recalls the morphological resemblance with the extant species Lepidopleurus cajetanus.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	materials_examined	Material examined. Middle Miocene: Central Paratethys: Austria: Pötzleinsdorf, type material. Upper Miocene: France, Ligerian Basin (Tortonian): Anjou: Saint-Clément-de-la-Place: type material plus 4 valves (BD 312). Italy, Po Basin (Tortonian): Borelli: 2 valves (BD 313), Montegibbio: 1 valve (BD 314), Rio di Bocca d’Asino: 3 valves (BD 315). Lower Pliocene: Italy: Liguria: Borzoli: 26 valves (BD 316), Bussana: 17 valves (BD 317), Genova Sestri: 2 valves (BD 318), Rio S. Antonino: 2 valves (BD 319). Pliocene: Italy: Tuscany: Pietrafitta: 1 valve (BD 320), Serre di Rapolano: 10 valves (BD 321), Stroncoli: 4 valves (BD 322); Emilia-Romagna: Bacedasco: 3 valves (BD 323), Vernasca: 1 valve (BD 324). Pleistocene: France: Corsica: Patrimonio: 2 valves (BD 325). Italy: Emilia-Romagna: Torrente Stirone: 4 valves (BD 326); Puglia: Punta Penne: 2 valves (BD 327); Calabria: Archi San Francesco: 4 valves (BD 328), Le Castella: 2 valves (BD 329), Pecoraro: 1 valve (BD 330), Pezzo: type material plus 4 valves (BD 331). Maximum width of the valves: 9.5 mm (intermediate).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	diagnosis	Diagnosis. Head and tail valves unknown. Intermediate valves solid, broadly rectangular, rounded, highly elevated, apex not evident, lateral areas triangular, originating from posterior margin, very angled. Tegmentum coarse, very variable, LA sculptured with strong, concentric, terraced ribs, CA with longitudinal chains of granules, somewhat branching or anastomosing. Granules of irregular shape, elevated, united with each other, with one subcentral megalaesthete and some micraesthetes. Articulamentum without insertion laminae, apophyses small, triangular.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	description	Description. Head and tail valves unknown. Intermediate valves solid, broadly rectangular, with a very variable width / length ratio, rounded in anterior profile, highly elevated (H / W = 0.40 – 0.64), anterior margin straight or slightly convex between apophyses, lateral margins slightly rounded, posterior margin very irregular, convex, apex not evident, lateral areas triangular, more irregular and sometimes slightly curved, originating from posterior margin, forming an angle from 50 - 60 ° upwards, up to almost perpendicular. Tegmentum coarse, very variable, LA sculptured with strong, concentric, terraced ribs, more or less spaced from each other, CA with longitudinal chains of granules, somewhat branching or anastomosing, transversally intersected by much thinner cords that give a pitted appearance. Granules of irregular shape, from roundish to polygonal, elevated, united with each other, with one subcentral megalaesthete and some micraesthetes. Articulamentum without insertion laminae, apophyses small, triangular, divided by a wide jugal sinus.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	discussion	Remarks. Lepidopleurus pseudocajetanus sp. nov. is characterized by intermediate valves [attributed to L. cajetanus (Poli, 1791) by previous authors] showing the starting point of LA with the concentric terraced ribs neighbouring the lateral margin (not near the apex), much more angled at the posterior margin, from 50 - 60 ° upwards up to almost perpendicular, rounded in anterior profile but more elevated (H / W = 0.40 - 0.64), which give the valves a very different look to that of L. cajetanus. The posterior margin is also very irregular, not straight but convex, as if there were real “ shortcomings ” (Figs 6 A, 6 D, 6 I), which however cannot be detected by an examination of the articulamentum, and with no trace of the apex. Similarly, the apical area is thicker, it does not tend to thin towards the ends of the posterior margin as in L. cajetanus, but projects towards the end of LA (Figs 6 B, 6 G). Some of these valves have already been photographed and published as L. cajetanus (Dell’Angelo & Palazzi 1989; Dell’Angelo et al. 2013, 2015 a, 2021 b). Intermediate plates of both species (Lepidopleurus cajetanus and L. pseudocajetanus sp. nov.) co-occur in some of the locations examined, including the Paratethys. Comparisons. See Tab. 1 for a comparison with the Lepidopleurus spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF24E0F0FADFB386BAF9660.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria; Pötzleinsdorf (this study). Upper Miocene: northeastern Atlantic: France: Saint-Clément-de-la-Place, (this study); Proto-Mediterranean Sea: Po Basin, N Italy: Borelli, Montegibbio, Rio di Bocca d’Asino (this study). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Genova Sestri, Rio S. Antonino (this study). Pliocene: central Mediterranean, Italy: Bacedasco, Pietrafitta, Serre di Rapolano, Stroncoli, Vernasca (this study). Pleistocene: central Mediterranean, France: Corsica: Patrimonio (this study); Italy: Archi S. Francesco, Le Castella, Pecoraro, Pezzo, Punta Penne, Torrente Stirone (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	description	Fig. 7	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	materials_examined	Type material. Holotype: MSNG 62625, intermediate valve, width 8.6 mm (Figs 7 E – F). Paratype 1: MNHN. F. A 98469, intermediate valve, width 9 mm (Figs 7 G – H); Paratype 2: SMF 380818, tail valve, width 6.6 mm (Figs 7 I – J); Paratype 3: MSNG 62626, tail valve, width 4.3 mm (Figs 7 K – L); Paratype 4: MSNG 62627, head valve, width 6 mm, (Figs 7 A – D). Type locality. Salice (Sicily, Italy). Type stage. Lower Pleistocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	etymology	Etymology. The name honors Agatino Reitano (Messina, Italy), for his contribution to the study and research of fossils from the Sicily.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	materials_examined	Material examined. Pleistocene: Italy: Sicily: Salice: type material plus 10 valves (BD 332), Furnari: 1 valve (BD 257). Maximum width of the valves: 6 / 9 / 6.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	diagnosis	Diagnosis. Valves solid, head valve semicircular, intermediate valves broadly rectangular, rounded, elevated, apex inconspicuous, tail valve elliptical, mucro flat, subcentral. Tegmentum coarse, sculptured with roundish granules irregularly disposed and concentric and elevated ribs in HV, LA, PMA, with many longitudinal chains of elongate granules and inconspicuous concentric ribs in CA, AMA. Articulamentum without insertion laminae, apophyses wide.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	description	Description. Valves solid. Head valve semicircular, posterior margin almost straight, front slope straight. Intermediate valves wide, broadly rectangular, length about half the width, rounded in anterior profile, elevated (H / W = 0.43 – 0.45), anterior margin slightly convex, side margins very rounded, posterior margins straight with apex inconspicuous, lateral areas strongly raised. Tail valve elliptical (W / L = 1.63 – 1.74), anterior margin convex, mucro flat, in central position, antemucronal slope slightly convex, postmucronal slope almost straight. Tegmentum coarse. HV, LA and PMA sculptured with numerous and well separated each other roundish granules, irregularly disposed, with presence of concentric and elevated ribs, 2 - 3 stronger in LA, someone more and less marked in HV and PMA. CA and AMA sculptured with many longitudinal chains of elongate granules, finer in jugal area, slightly oblique near side margins, with inconspicuous, barely visible, concentric ribs. Articulamentum without insertion laminae, apophyses wide.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	discussion	Remarks. Coherently with most Lepidopleuru s spp., the valves of this taxon are rather robust, although not perfectly preserved so that we have no complete intermediate valves. The sculpture is characteristic, the longitudinal chains of granules on the intermediate valves are fairly regular, the intercostal spaces are narrow and smooth. The granules are irregularly disposed in LA and PMA, not arranged in radial chains. Comparisons. The sculpture of Lepidopleurus reitanoi sp. nov. is different from that of the other species of Lepidopleurus here discussed. The longitudinal chains of granules of CA and AMA are broadly regular, not more or less branching or anastomosing as in L. cajetanus (Poli, 1791), L. benoisti (de Rochebrune, 1882), L. pseudobenoisti Dell’Angelo et al., 2018 b and L. virgifer (Sandberger, 1859). Further, the sculpture of LA with roundish granules irregularly disposed and some concentric ribs is different from that of L. cajetanus, L. benoisti, L. pseudobenoisti and L. virgifer, all showing irregular radial chains of granules, sometimes branching or anastomosing, intersected by concentric, more or less terraced ribs. Lepidopleurus gallicus Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 differs from Lepidopleurus reitanoi sp. nov. by the more numerous longitudinal chains of granules on AMA (less than half in L. reitanoi sp. nov.), and the radial chains of granules on PMA (vs. granules irregularly disposed in L. reitanoi sp. nov.).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFF04E300FADFC3C6E9393E9.taxon	distribution	Distribution. Lower Pleistocene: central Mediterranean, S. Italy: Salice, Furnari (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	description	Fig. 8	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	materials_examined	Type material. Syntypes: NHMW 1862 / 0012 / 0047, 2 intermediate valves; NHMW 1863 / 0017 / 0062, 3 valves (Figs 8 B – D). Type locality. Gienberg (Germany). Type stage. Oligocene (Rupelian), Alzey Formation. Material examined. Lower Oligocene: Germany, Mainz Basin: type material, plus Gienberg: 9 valves (BD 333, NHMW 1868 / 0001 / 0776, NHMW 1868 / 0001 / 0777, Figs 8 I – J); Steigerberg: 5 valves (BD 334, Figs 8 A, 8 EH, 8 K – L). Maximum width of the valves: 6.1 / 9.7 / 6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	description	Description. Valves solid. Head valve large, semicircular, widely open posteriorly. Intermediate valves wide, broadly rectangular, width more than three times the length (W / L = 3.13), rounded in anterior profile, moderately elevated (H / W = 0.39), anterior and posterior margins straight, side margins rounded, apex inconspicuous, lateral areas strongly raised. Tail valve semicircular (W / L = 1.72 – 1.87), elevated, anterior margin from almost straight to slightly convex in jugal area, mucro pronounced, in anterior-subcentral position, antemucronal slope convex, postmucronal slope concave just behind mucro. Tegmentum very coarse, with a granulated-nodulose sculpture. HV, LA and PMA sculptured with numerous and strongly irregular branching or anastomosing radial chains of roundish granules, coarser in LA, more variable in PMA, intersected by some more or less evident concentric, terraced ribs. CA and AMA sculptured with longitudinal chains of roundish granules, united with each other, diameter up to 100 µm, more irregular and rougher, shaped by groups of granules branching longitudinally many times in CA, more regular in AMA with longitudinal chains of granules finer and less subject to split. Each granule with a central megalaesthete and a series of micraesthetes (normally 6 – 8) spaced along margin. Articulamentum without insertion laminae, apophyses narrow, triangular in intermediate valves, larger and rounded in tail valves, widely projecting.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	discussion	Remarks. The study of the syntype material hosted at NHMW (Dell’Angelo et al. 2019 b) revelead a certain variability in intermediate and tail valves (head valves were not present), not so obvious in the original description. Sandberger described and figured a tail valve “ with an almost exactly hemicircular margin ”, while the syntypes show an anterior margin from almost straight to slightly convex, and a more variable position of the mucro, not in such an anterior position as mentioned in the original description. The antemucronal and postmucronal slopes of the tail valve (never described and illustrated by Sandberger or Janssen 1978) also show certain variability, mainly in the more or less accentuated concavity of the postmucronal slope (Dell’Angelo et al. 2019 b: figs 2 K, 2 O). Šulc (1934) considered large tail valves (width of 18 mm) of Lepidopleurus decoratus Reuss, 1860 from the Middle Miocene of Pötzleinsdorf (Austria) to be close to L. virgifer (see above, check L. benoisti). Comparisons. Lepidopleurus virgifer (Sandberger, 1859) differs from the other Oligocene species largely by the very coarse granulated-nodulose sculpture, which is even visible in juvenile segments, especially in the central area of intermediate valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	distribution	Distribution. Lower Oligocene: North Europe, Germany: Böseberg, Gienberg, Glimmerode, Steigerberg, Zeilstück, Würzmühle (Janssen 1978; Hocht 1986; Dell’Angelo et al. 2019 b; this study). Genus Leptochiton Gray, 1847	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	type_taxon	Type species. Chiton cinereus Montagu, 1803 (misapplication of name), by subsequent designation (Gray 1847 a), not Linnaeus, 1767 (= Chiton asellus Gmelin, 1791).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	distribution	Distribution. Leptochiton is known from the Triassic to the Recent, with a living worldwide distribution, most species from the northeastern Atlantic and the Mediterranean Sea. Fossils determined as Leptochiton have been found worldwide and Leptochiton is one of the most ancient genera of “ modern ” polyplacophorans, dating back to the Triassic (Laghi 2005), the Jurassic of France, Germany, Poland, Russia and Siberia (Sirenko 2013), and possibly even to the lower Carboniferous (Sirenko 2013). The fossil record includes the Paleocene of Denmark (Sigwart et al. 2006), the Eocene of Europe, Australia (Dell’Angelo et al. 2011), Antarctica (Lopez Cabrera & Olivero 2011), the upper Eocene-lower Oligocene of Washington, U. S. A. (Dell’Angelo et al. 2011), the Oligocene of Europe (France and Germany: Dell’Angelo et al. 2011, 2018 a) and New Zealand (Lee et al. 2014; Wu & Lee 2024), the Miocene of the Paratethys (Šulc 1934; Bałuk 1971, 1984), the Miocene-Pleistocene of the Mediterranean Basin (Dell’Angelo et al. 2013, 2016), Australia (Cotton 1964) and New Zealand (Beu & Maxwell 1990; Sutherland et al. 1995), the Pliocene-Pleistocene of California (Vendrasco et al. 2012), and the Holocene of Japan (Kuroda et al. 1980).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFCF4E3C0FADF8B46E4297CC.taxon	discussion	Remarks. Sigwart et al. (2011) attempted to elucidate the relationships of the earliest diverging living chitons of the order Lepidopleurida using molecular phylogenetic methods. A result of this work found that the large cosmopolitan genus Leptochiton is not monophyletic and is restricted to the North Atlantic and the Mediterranean Sea. However, there are several groups of deep-water leptochitonids with complexes of similar morphological features which inhabit low latitudes in the Pacific, Indian and Atlantic Oceans (Sirenko 2015). The genus Leptochiton is characterized by the lack of insertion laminae, the apophyses small and neatly separate, and the tegmentum uniformly granulated. For a better identification and differentiation, the examined species are reported, in the tables that summarize the main diagnostic characters, in three groups which highlight the different tegmentum sculpture: Leptochiton cancellatus group: HV, LA and PMA with granules arranged in radial series; CA and AMA with granules arranged in longitudinal series. The 16 species belonging to this group are most uncommon or rare, fragile and seldom found in good conditions, so they are not easy to determine. The main morphological characters of the species of Leptochiton cancellatus group considered in the present study are reported in Tabs 2 and 3. ...... continued on the next page ...... continued on the next page ...... continued on the next page	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC34E3D0FADFC646F0B9174.taxon	description	Fig. 9	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC34E3D0FADFC646F0B9174.taxon	materials_examined	Type material. Holotype MZB 7074, intermediate valve, width 3.6 mm (Figs 9 A – E); Paratype BD 3828, intermediate valve, width 1.6 mm. Type locality. Pezzo, near Villa S. Giovanni (Calabria, Italy). Type stage. Upper Pleistocene, but possibly reworked from contiguous deposits of the Lower Pleistocene of bathyal facies. Material examined. Upper Pleistocene: Pezzo (Italy): type material and another intermediate valve (BD 335, Figs 9 F – H), width 3.5 mm. Maximum width of the valves: 3.6 mm (intermediate).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC34E3D0FADFC646F0B9174.taxon	description	Description. Head and tail valves unknown. Intermediate valves broadly rectangular, semicarinate in anterior profile forming angle of ca 80 °, highly elevated (H / W = 0.60 – 0.68), anterior margin concave in wide jugal area, antero-lateral corners obliquely truncated, side margins slightly rounded, posterior margin almost straight at both sides of small apex, lateral areas very raised. Tegrnentum rough, sculptured with roundish-polygonal, separated granules, irregularly arranged, less evident in JA, forming longitudinal irregular ribs in hind zone of CA, for about half valve; LA furrowed by strong concentric growth lines in continuity with those of CA. Articulamentum without insertion laminae, apophyses wide, semicircular.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC34E3D0FADFC646F0B9174.taxon	discussion	Remarks. This species has been found only from the type locality, and the head and tail valves are not known. At the time of the original description, only the two valves designated as holotype and paratype were known. The holotype is illustrated here for the first time (only by drawings in the original description). The paratype is smaller and differs from the holotype for a lowest elevation, a less pronounced jugal angle, and in lacking the longitudinal irregular ribs in the hind zone of CA. Subsequently, another intermediate valve was found, shown here (Figs 9 F – G), matching to the characters of Leptochiton abacinus (Dell’Angelo & Palazzi, 1989). The anterior profile seems a little more carinated (Fig. 9 G), the height / width ratio is 0.60 (vs. 0.68 of the holotype), and the angle of anterior profile is 80 °, as in the holotype. Comparisons. Leptochiton abacinus differs from all Leptochiton species discussed here, mainly because of its considerable elevation (H / W = 0.60 – 0.68) and for the peculiar sculpture with irregularly arranged granules, forming longitudinal irregular ribs only in the hind zone of PA for about half valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC34E3D0FADFC646F0B9174.taxon	distribution	Distribution. Pleistocene: central Mediterranean, S. Italy: Pezzo (Dell’Angelo & Palazzi 1989; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC24E3F0FADFB096A949458.taxon	description	Fig. 10	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC24E3F0FADFB096A949458.taxon	materials_examined	Type material. Unknown, possibly in École des Mines, Paris (fide Kaas & Van Belle 1985 a: 44). Type locality. Algeciras (Spain). Material examined. Miocene (Tortonian): France: Saint-Clément-de-la-Place: 20 valves (MNHN. F. A 67062 – A 67066, Figs 10 A – E, NHMW 2017 / 0108 / 0007, RGM. 1008428, BD 133); Renauleau: 1 valve (BD 134). Pliocene: Spain: Estepona: 3 valves (BD 336). Italy: Tuscany: Colle Val d’Elsa: 3 valves (BD 337, Figs 10 F – H), Pietrafitta Melograni: 7 valves (BD 338); Sicily: Altavilla: 1 valve (BD 339). Upper Pliocene to lower Pleistocene: France: Bosq d’Aubigny: 7 valves (RGM. 1310182, RGM. 1310183). Pleistocene: Italy: Calabria: Musalà: 8 valves (BD 340), Pecoraro: 3 valves (BD 341), Pezzo: 8 valves (BD 342), Sicily: Ficarazzi: 54 valves (BD 343). Maximum width of the valves: 5 / 5 / 8.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC24E3F0FADFB096A949458.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, notched in the middle. Intermediate valves broadly rectangular (W / L = 2.70 – 3.40), moderately elevated (H / W = 0.22 – 0.40), rounded in anterior profile, anterior margin straight, side margins rounded, posterior margin straight, apex not indicated, lateral areas hardly raised, though clearly indicated. Tail valve semicircular (W / L = 1.66 – 1.91), front margin convex, mucro central, somewhat swollen but not prominent, antemucronal slope slightly convex, postmucronal slope straight. Tegmentum rough, space between striae of granules reduced, generally with 4 - 5 obvious growth wrinkles encircling the shell completely. HV, LA and PMA sculptured with small roundish, united granules arranged in radiating series (HV 100 or more, LA 14 – 16). CA and AMA sculptured with roundish, united granules arranged in longitudinal series (CA 65 – 70), the series becoming slightly converging towards the outer margins. Each granule in CA and AMA with a central megalaesthete and a series of micraesthetes (normally 5 – 6, up to 8 or more) along edge. Articulamentum lacking insertion laminae, apophyses small, broadly triangular, widely separated, evenly rounded in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC24E3F0FADFB096A949458.taxon	discussion	Remarks. Fossil records for this species are rather scant and historical records require confirmation (Dell’Angelo & Palazzi 1989; Dell’Angelo & Smriglio 1999). Dell’Angelo & Palazzi (1989) considered Lepidopleurus maguntiacus de Rochebrune, 1882 from the middle Oligocene of western Germany as a synonym of the present species. The two species undoubtedly have remarkable similarities, discussed by Dell’Angelo & Palazzi (1989), but they differ in a series of characters, highlighted in the discussion of L. maguntiacus (see below), and in the different stratigraphic distribution, limited to the lower Oligocene for L. maguntiacus, not exceeding the Miocene (Tortonian) for L. algesirensis (Capellini, 1859). There is a great variability in the material examined, especially for the shape of intermediate (W / L = 2.70 – 3.40, H / W = 0.22 – 0.40) and tail (W / L = 1.66 – 1.91) valves. Comparisons. See Tab. 2 for a comparison with the Leptochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC24E3F0FADFB096A949458.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Anjou, France: Saint-Clément-de-la-Place, Renauleau (Dell’Angelo et al. 2018 b). Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Altavilla, Colle Val d’Elsa, Pietrafitta Melograni (Dell’Angelo & Palazzi 1989; Dell’Angelo et al. 2001 a; Chirli 2004; Dell’Angelo et al. 2012). Upper Pliocene to upper Pleistocene: northeastern Atlantic, Ligerian Basin, France: Bosq d’Aubigny (Dell’Angelo et al. 2018 b). Pleistocene: central Mediterranean, S. Italy: Ficarazzi, Musalà (Dell’Angelo & Palazzi 1989), Pecoraro, Pezzo (this study). Recent: eastern Atlantic Ocean: from Portugal and Berlengas Arch. (Pisani Burnay 1986; Consolado Macedo et al. 1999) south to Morocco and Senegal (Leloup 1968), Canary Islands (Leloup 1968; Hernández & Rolán 2011), Madeira and Selvagens Arch. (Segers et al. 2009; Kaas 1991); Mediterranean Sea: Italy (Dell’Angelo & Smriglio 1999), Aegean Sea and Levantine Basin (Koukouras & Karachle 2005), Tunisia: Gulf of Gabes (Cecalupo et al. 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC04E200FADFE246B699660.taxon	description	Fig. 11	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC04E200FADFE246B699660.taxon	materials_examined	Type material. Holotype MZUB 60418, intermediate valve, width 3.4 mm, Figs 11 A – D. Paratypes: MZUB 60419 intermediate valve, width 2.9 mm; MZUB 60420, tail valve, width 3.1 mm, Figs 11 I – L. Type locality. Southwestern Adriatic Sea, Apulian margin, off Bari (Italy), cruise SE 06 - 50. Type stage. Upper Pleistocene submerged deposits, probably late glacial epoch. Material examined. Type material, plus: SE 06 - 10: 1 valve; SE 06 - 35: 4 valves; SE 06 - 40: 3 valves, Figs 11 EH; SE 06 - 48: 2 valves; SE 06 - 50: 55 valves. Maximum width of the valves: -- / 3.4 / 3.1 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC04E200FADFE246B699660.taxon	description	Description. Head valve unknown. Intermediate valves broadly rectangular (W / L = 1.45 – 2.03), rounded in anterior profile, elevated (H / W = 0.50 – 0.66), anterior margin straight, side margins rounded, posterior margin straight, apex inconspicuous, lateral areas hardly or not raised. Tail valve semicircular (W / L = 1.48 – 1.61), anterior margin almost straight or slightly convex, mucro not prominent, in slightly anterior position, antemucronal slope slightly convex, postmucronal slope almost straight. Tegmentum rough, space between striae of granules reduced. LA and PMA with granules more rectangular, randomly disposed along concentric lines, with numerous and well-marked growth lines; granules with a maximum width up to 60 µm and presence of 3 – 4 aesthetes more or less aligned, with pores of same width. CA and AMA with well raised, rather thick but distinctly separated roundish to oval granules, extended with 2 – 3 small longitudinal varices, forming 50 – 55 longitudinal series with a regular quincuncial pattern displayed by granules of neighboring rows; granules with a maximum width up to 70 µm and the presence of one subcentral aesthete and up to 6 aesthetes irregularly disposed, with pores of same width. Articulamentum without insertion laminae, with apophyses small, sharply triangular, widely separated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC04E200FADFE246B699660.taxon	discussion	Remarks. The fossil record of Leptochiton antondohrni Taviani, Sosso & Dell’Angelo, 2023 refers to the late Pleistocene (presumably last glacial) of the Adriatic Sea, although we cannot completely exclude that is still alive in the Mediterranean Sea (Taviani et al. 2023). The material studied is quite well preserved, and the intermediate valves show some variability. Two types of intermediate valves are detectable, some higher (Figs 11 A – D, H / W = 0.66) and slightly larger (W / L = 1.45 – 1.60), others a little less high (Figs 11 E – H, H / W = 0.50 – 0.55) and a little smaller (W / L = 1.91 – 2.03). The structure of the granules also shows slight differences, the granules in CA are more oval and up to 70 µm in size in the higher valves, more rounded and slightly smaller up to 60 µm in the lower ones. Comparisons. Leptochiton antondohrni differs from L. prudenzae sp. nov. by the valves slightly larger, the intermediate valves more elevated (W / L = 0.50 – 0.66 vs. 0.33 – 0.37 in L. prudenzae), and the larger granules (up to 60 – 70 µm vs. 38 µm in L. prudenzae).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFC04E200FADFE246B699660.taxon	distribution	Distribution. Pleistocene, presumably last glacial: central Mediterranean, Italy: Adriatic Sea, offshore Bari (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDF4E220FADFC3C69AB9458.taxon	description	Fig. 12	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDF4E220FADFC3C69AB9458.taxon	materials_examined	Type material. Holotype: MSNP, a specimen collected on Posidonia at a depth of 10 m (Figs 12 A – D). Paratypes: 1 intermediate valve from Porto Cesareo (MZB); specimen from Capraia Island (AL); valves from Elba Island, Vendicari and Creta (Greece) (BD 2921, BD 3031, BD 3388). Type locality. Laghi Alimini (Lecce, Italy). Material examined. Lower Pliocene: Italy: Liguria: Borzoli: 2 valves (BD 344), Bussana: 2 valves (BD 345), Genova Sestri: 1 valve (MZB 45703, Figs 12 E – H), Rio S. Antonino: 5 valves (MP), Rio Torsero: 1 valve (BD 346). Pliocene: Italy: Tuscany: Pietrafitta: 3 valves (BD 347), Pietrafitta Melograni: 5 valves (BD 348), Poggio alla Fame: 3 valves (BD 349, Figs 12 I – J), Serre di Rapolano: 2 valves (BD 350). Recent: Italy: type material, plus Favignana Island: valves from cruise CS 96 - 88 (MZB, Figs 12 K – L). Maximum width of the valves: -- / 2.5 / 2.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDF4E220FADFC3C69AB9458.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, notched in middle, front slope straight. lntermediate valves broadly rectangular (W / L = 2.28 – 2.54), rounded to semicarinate in anterior profile, moderately elevated (H / W = 0.38 – 0.45), anterior margin straight to slightly convex, side margins rounded, posterior margin straight, apex inconspicuous, lateral areas moderately raised. Tail valve elliptical, length 2 / 3 of width (W / L = 1.40), anterior margin consisting of three segments, mucro subcentral, rather prominent, antemucronal slope straight to slightly convex, postmucronal slope concave directly behind mucro. Tegmentum smooth, space between striae of granules large, up to twice width of the series of granules, growth lines barely noticeable. HV, LA and PMA sculptured with minute roundish united granules partially overlapping, arranged in radial striae, often bifurcating (HV 60 dentating outer margin, about 20 granules per series, LA 6 – 8, PA 50). CA and AMA sculptured with minute roundish united granules partially overlapping, arranged in longitudinal striae (CA 30) LA and CA connected in very regular and characteristic way. Each granule with a central megalaesthete and many micraesthetes (up to 15 or more) irregularly spaced all along, except on upper side. Articulamentum lacking insertion laminae, apophyses small, triangular, widely separated by straight sinus, about 1 / 3 width of valve, apical area expanded, quadrangular, with upper edge bisinuated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDF4E220FADFC3C69AB9458.taxon	discussion	Remarks. This elusive species has been described based on three living individuals and a few loose Recent valves collected at Laghi Alimini (Lecce), Capraia Island (Tuscan Archipelago), and Punta Prosciutto (Taranto). The fossil record of Leptochiton bedullii Dell’Angelo & Palazzi, 1986 is limited to the Lower Pliocene of Liguria (Dell’Angelo et al. 2013) and Tuscany (Dell’Angelo et al. 2001 a; Chirli et al. 2004). Dell’Angelo & Palazzi (1989) considered Leptochiton bedullii as a synonym of L. boettgeri (Šulc, 1934), a species known from the Miocene of Kostej (Romania) only for two valves (one head and one tail). We consider these two taxa as distinct species (see discussion below regarding Leptochiton boettgeri). Some reports of Mediterranean specimens considered as Leptochiton boettgeri on the basis of this synonymy (e. g., Öztürk et al. 2014) must therefore be correctly attributed to Leptochiton bedullii. A single intermediate valve from Altavilla, illustrated by Dell’Angelo & Palazzi (1989: pl. 20, fig. 1) as Lepidopleurus (Leptochiton) boettgeri (considered by these authors conspecific with Leptochiton bedullii), was left undetermined at species level by Dell’Angelo et al. (2012); consequently, L. bedullii is excluded from the chiton fauna of the Pliocene of Altavilla. Comparisons. See Tab. 2 for a comparison with the Leptochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDF4E220FADFC3C69AB9458.taxon	distribution	Distribution. Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Rio S. Antonino, Rio Torsero, Sestri Ponente (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b). Pliocene: central Mediterranean, Italy: Tuscany: Pietrafitta, Serre di Rapolano (Dell’Angelo et al. 2001 a; Chirli 2004), Poggio alla Fame (this study). Recent: Mediterranean Sea: Italy (Dell’Angelo & Smriglio 1999), Croatia: Mljet National Park (Dell’Angelo & Zavodnik 2004), Malta, Greece and Aegean Sea Islands (Strack 1988), Turkey (Ozturk et al. 2014), Lebanon (Crocetta et al. 2014), Tunisia (Kaas 1989; Cecalupo et al. 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDD4E240FADFE246BD99380.taxon	description	Fig. 13	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDD4E240FADFE246BD99380.taxon	materials_examined	Type material. Unknown, probably lost (fide Kaas & Van Belle 1985 a: 43). Type locality. P ossibly the coast of Great Britain, probably Oban, Scotland (fide Kaas & Van Belle 1985 a: 43). Material examined. Middle Miocene: Eastern Paratethys: Ukraine: Varovtsi: 7 valves (BD 351, Figs 13 M – P), Horodok: 12 valves (BD 352, Figs 13 Q – T). Upper Miocene: Italy: Po Basin: Montegibbio: 2 valves (BD 353, MZB 32046), Rio di Bocca d’Asino: 8 valves (BD 354, PG, MZB 32010 – 32011, Figs 13 I – K), Villa Monti: 1 valve (MZB 32045). Lower Pliocene: Italy: Liguria: Borzoli: 42 valves (BD 355), Bussana: 1 valve (BD 356), Caranchi: 2 valves (MP), Genova Sestri: 1 valve (BD 357), Rio S. Antonino: 16 valves (MP, MZB 45699 – 45700, Fig. 13 L), Rio Torsero: 1 valve (BD 358). Pliocene: Portugal: Vale de Freixo: 265 valves (BD 237, GeoFCUL VFX. 03.339, GeoFCUL VFX. 03.346, GeoFCUL VFX. 03.349, Figs 13 E – G, MNHN. F. A 81981, RGM. 1363997 – 1363998). Italy: Tuscany: Cetona: 1 valve (BD 359), Orciano Pisano: 2 valves (BD 360), Pietrafitta: 4 valves (BD 361), Pietrafitta Melograni: 4 valves (BD 362), Poggibonsi: 1 valve (BD 363), Poggio alla Fame: 3 valves (BD 364, Figs 13 A – D), Serre di Rapolano: 1 valve (BD 365); Emilia-Romagna: Cava di Campore: 42 valves (BD 366), Lugagnano: 4 valves (BD 367), Rio Stramonte: 2 valves (BD 368). Upper Pliocene-Pleistocene: Italy: Sicily: Altavilla: 8 valves (AG, AR, BD 369). Pleistocene: Italy: Tuscany: Caletta: 2 valves (BD 370); Calabria: Carrabbati: 4 valves (BD 371), Torrente Boscaino: 1 valve (BD 372), Vallone Catrica: 1 valve (BD 373); Sicily: Salice: 5 valves (BD 374, Fig. 13 H), Sciacca: 1 valve (BD 375). Maximum width of the valves: 2.5 / 3.4 / 3.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDD4E240FADFE246BD99380.taxon	description	Description. Valves thin and fragile. Head valve less than semicircular, posterior margin widely V-shaped, front slope straight. Intermediate valves broadly rectangular (W / L = 2.38 – 3.23), moderately elevated (H / W = 0.30 – 0.44), rounded in anterior profile, anterior margin slightly convex, side margins slightly rounded, posterior margin about straight, apex not indicated, lateral areas raised, mostly with a few more or less prominent growth ridges. Tail valve semicircular (W / L = 1.55 – 1.90), anterior margin straight to slightly convex, mucro subcentral, swollen but not prominent, antemucronal slope slightly convex, postmucronal slope slightly concave. Tegmentum rough, space between striae of granules reduced, growth lines barely noticeable. HV, LA and PMA sculptured with dense oval granules united, arranged in radial series (HV ca. 50 – 60, LA ca. 10). CA and AMA sculptured with roundish granules united, partially overlapping, arranged in longitudinal series (CA ca. 60). Each granule with a central megalaesthete and a series of micraesthetes (normally 6 – 7) rather regularly spaced all along, except on upper side. Articulamentum without insertion laminae, apophyses small, well separated by a slightly concave jugal sinus, triangular but tending to be trapezoidal in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDD4E240FADFE246BD99380.taxon	discussion	Remarks. Most of the records of small Mediterranean Leptochiton collected at depth between 30 and 100 m have been attributed in the past to L. cancellatus (Dell’Angelo & Palazzi 1986), whereas this species is instead uncommon in the Mediterranean, being found mainly in deep-water detrital bottoms (Dell’Angelo & Smriglio 1999). The fossil record of Leptochiton cancellatus is scant; the species is known from a few sites from Miocene, more common from the Pliocene, rarely recorded from the Pleistocene of Italy, and from the North Europe (Sweden and Norway). The recent findings of Leptochiton cancellatus from the Paratethys (Studencka & Dulai 2010; this study, Figs 13 M – T) considerably extend the stratigraphic and geographic distribution of the species.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDD4E240FADFE246BD99380.taxon	materials_examined	There is a great variability in the material examined, especially for the shape of intermediate valves, from W / L = 2.38 – 2.50 (Recent) to 2.60 – 2.80 (Paratethys) and 3.05 – 3.23 (Pliocene of Italy and Portugal). Comparisons. See Tab. 2 for a comparison with the Leptochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDD4E240FADFE246BD99380.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Poland: Gieraszowice, Rybnitsa, Węglinek (Macioszczyk 1988; Studencka & Studencki 1988); Eastern Paratethys: Ukraine: Horodok, Varovtsi (Studencka & Dulai 2010; this study). Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, North Italy: Montegibbio, Rio di Bocca d’Asino, Villa Monti (Dell’Angelo & Palazzi 1989; Dell’Angelo et al. 2015 a). Lower Pliocene: North Europe, Belgium: Kallo (Marquet 1984, 2002); northeastern Atlantic: U. K. (Reid 1890); central Mediterranean, Italy: Liguria: Borzoli, Caranchi, Rio S. Antonino, Rio Torsero, Sestri Ponente (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b); Pliocene: northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo & Silva 2003; Dell’Angelo et al. 2022) central Mediterranean, Italy: Emilia-Romagna: Cava di Campore, Lugagnano Val d’Arda, Rio Stramonte (this study); Tuscany: many localities (Dell’Angelo et al. 2001 a; this study): Sicilia: Altavilla Milicia (Dell’Angelo et al. 2012). Pleistocene: North Atlantic: Sweden and Norway (Brogger 1901; Antevs 1917), Netherlands (Strack 2010); central Mediterranean, S. Italy: Carrabbati, Salice, Sciacca, Torrente Boscaino, Vallone Catrica (this study). Recent: Atlantic Ocean: from U. K. and Ireland (McKay & Smith 1979; Light & Baxter 1990) to the coast of France (Kaas 1979; Dell’Angelo & Palazzi 1986), Spain (Borja 1987; Rolan Mosquera et al. 1990; Urgorri et al. 2017), Portugal (Consolado Macedo et al. 1999) and Madeira Arch. (Segers et al. 2009). Mediterranean Sea: Spain (Moreno & Gofas 2011); Italy: Gulf of Venezia (Cesari 1987), Banco Amendolara (Panetta et al. 1985), Strait of Messina (Giacobbe & Renda 2018), Strait of Sicily (Dell’Angelo et al. 1998 a), and many other localities (Dell’Angelo & Smriglio 1999); Malta (Mifsud et al. 1990); Greece and Aegean Sea Islands (Strack 1990; Zenetos & Van Aartsen 1995; Koukouras & Karachle 2005); Turkey (Ozturk et al. 2014); Tunisia (Kaas 1989; Cecalupo et al. 2008); Israel (Barash & Danin 1977); Marmara Sea (Öztürk et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	description	Fig. 14	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	materials_examined	Type material. Holotype: MSNG 62628, tail valve, width 1.3 mm (Figs 14 A – C). Paratype: MSNG 62629, tail valve, width 1 mm (Figs 14 D – H). Type locality. Archi (Calabria, Italy). Type stage. Lower Pleistocene (a bathyal paleoenvironment, 500 to 1000 m paleodepth: Di Geronimo et al. 1997).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	etymology	Etymology. The species is dedicated to Franco Corticelli (IMM-CNR Bologna) as a recognition of his valuable and skilful support to produce most of the S. E. M. images of this study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	materials_examined	Material examined. Italy: Archi: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	diagnosis	Diagnosis. Tail valve semicircular, mucro in slighty posterior position. Tegmentum rough, sculptured with elevated granules, irregularly arranged, with 2 – 3 stems in apical part, roundish in AMA, flattened oval, almost rectangular, in PMA, each granule with three aesthetes almost aligned. Articulamentum without insertion laminae, apophyses small, triangular.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	description	Description. Head and intermediate valves unknown. Tail valve semicircular, anterior margin straight, mucro in slightly posterior position, not prominent, antemucronal slope slightly convex, postmucronal slope slightly concave. Tegmentum rough, with some isolated stem irregularly present, sculptured with elevated granules, irregularly arranged, with 2 – 3 stems in apical part, roundish in AMA (diameter up to 40 µm), flattened oval, almost rectangular, in PMA (length up to 40 µm), showing a more or less concentric arrangement. Stems give striated aspect to the tegmentum, and these are interrupted by granules. Each granule with three aesthetes almost aligned, one central megalaesthete and two micraesthetes on side. Articulamentum without insertion laminae, apophyses small, triangular, widely separated by a large jugal sinus.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	discussion	Remarks. Although only two tail valves are available in the material studied, the good state of conservation and the particular sculpture of the tegmentum, different from that of other known living and fossil species, allow us to describe it as a new species. Comparisons. Leptochiton corticellii sp. nov. is very similar to L. pepezamorai Carmona Zalvide, Urgorri & García, 2004, a living species known from Galicia (NW Spain) and from the Mediterranean area (Tuscan Archipelago, S. Lucia Bank: Dell’Angelo et al. 2009), a species not treated here because not known as a fossil, from which it differs mainly by the different arrangement of granules on AMA, randomly in Leptochiton corticellii sp. nov., vs. in longitudinal series in L. pepezamorai.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFDB4E250FADF89C6EF49291.taxon	distribution	Distribution. Lower Pleistocene: central Mediterranean, S. Italy: Archi (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD94E270FADFF05680094C8.taxon	description	Fig. 15	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD94E270FADFF05680094C8.taxon	materials_examined	Type material. Holotype MZUB 60348, intermediate valve, width 3.5 mm (Figs 15 A – D). Paratype 1: MZUB 60349, head valve, width 2.1 mm (Figs 15 E – F); Paratype 2: MZUB 60350 tail valve, width 2.2 mm; Paratype 3: MNHN-IM- 2022 - 2408, intermediate valve, width 3 mm; Paratype 4: MNHN-IM- 2022 - 2409, tail valve, width 2.7 mm (Figs 15 G – H); Paratype 5: SMF 366408, intermediate valve, width 2.9 mm; Paratype 6: SMF 366409, tail valve, width 2.7 mm Type locality. Capraia Island-Capo Corso, sediments - 350 / 500 m by fishermen. Type stage. Pleistocene, presumably last glacial. Material examined. Pleistocene, presumably last glacial: Italy: Tuscany: Capraia Island-Capo Corso - 350 / 500 m: type material plus 100 + valves (BD 248). Maximum width of the valves: 2.7 / 3.5 / 3.0 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD94E270FADFF05680094C8.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.43 – 2.85), rounded in anterior profile, elevated (H / W = 0.44 – 0.51), anterior margin almost straight, side margins rounded, posterior margin straight, apex not evident, lateral areas not raised with faint concentric lines of growth. Tail valve semicircular, elevated, anterior margin slightly convex, mucro subcentral, not prominent, antemucronal slope convex, postmucronal slope slightly concave. Tegmentum rough, space between striae of granules very reduced. HV, LA, PMA sculptured with small elliptical granules (up to 40 µm) moderately raised, with 2 – 3 stems in apical part, randomly arranged, faint concentric lines of growth in LA, more evident in HV, PMA. CA, AMA sculptured with granules roundish and more compact (up to 55 – 60 µm) arranged in longitudinal series (CA 80, AMA 60), with a regular quincuncial pattern displayed by granules of neighboring rows. Each granule with a variable number of aesthetes, up to 6 in CA and AMA, and 3 aesthetes lined up along diameter in HV, LA and PMA, all aesthetes of same size. Articulamentum without insertion laminae, weakly developed, apophyses small, triangular, wide apart in intermediate valves, trapezoid in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD94E270FADFF05680094C8.taxon	discussion	Remarks. This material studied by Dell’Angelo & Giusti (2000), has been attributed to Leptochiton sarsi Kaas, 1981 (see below). Leptochiton freiwaldi Dell’Angelo, Sosso & Taviani, 2024 is presently known from submerged assemblages of putative glacial Pleistocene age trawled offshore Capraia Island and Capo Corso in the Tyrrhenian Sea at a depth of 350 / 500 m. Comparisons. Leptochiton freiwaldi differs from Leptochiton sarsi mainly by the sculpture, with longitudinal striae of granules in CA and AMA more numerous and much closer, unlike L. sarsi, which shows tegmentum rough, and space between striae of granules large. Also the shape of the intermediate valves is different, with valves about twice as wide (W / L = 1.90 – 2.12) in L. sarsi, more elongated (W / L = 2.43 – 2.85) in L. freiwaldi.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD94E270FADFF05680094C8.taxon	distribution	Distribution. Pleistocene: central Mediterranean, Italy: Capraia Island-Capo Corso (Dell’Angelo & Giusti 2000; Dell’Angelo et al. 2024; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD84E280FADFDD468CE916C.taxon	description	Fig. 16	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD84E280FADFDD468CE916C.taxon	description	Dell’Angelo et al. 2001 a, p. 147, fig. 7.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD84E280FADFDD468CE916C.taxon	materials_examined	Type material. Holotype IRSN IG- 26.356; Paratype VB 2907 a, a specimen disarticulated off Bagur (Gerona, Spain), - 250 m, Figs 16 C – H. Type locality. off Llansa (Gerona, Spain), - 200 m. Material examined: Pleistocene: Italy: Riparbella: 1 valve, width 2.5 mm (BD 376, Figs 16 A – B).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD84E280FADFDD468CE916C.taxon	description	Description. Head valve semicircular, posterior margin almost straight. Intermediate valves broadly rectangular, semicarinate in anterior profile, moderately elevated (H / W = 0.37), anterior margin straight between the apophyses, somewhat backwardly directed towards the sides, side margins evenly rounded, posterior margin practically straight, apex inconspicuous, lateral areas hardly perceptible. Tail valve a little less than semicircular, width at least twice the length, mucro placed anteriorly and not prominent, postmucronal slope straight. Tegmentum rough, space between striae of granules large. HV, LA, PMA sculptured with roundish, clearly separated granules arranged in quincuncial pattern, giving the impression of radial rows near outer sides. CA, AMA sculptured with roundish granules with 2 large stems in apical part, arranged in longitudinal series (CA ca 40). Each granule with 3 aesthetes aligned, 1 megalaesthete subcentral and 2 micraesthetes at extremities, in CA. Articulamentum without insertion laminae, weakly developed, apophyses small, broadly triangular, trapezoid in tail valve, widely separated by a more or less flat sinus.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD84E280FADFDD468CE916C.taxon	discussion	Remarks. Leptochiton geronensis Kaas & Van Belle 1985 was described on the basis of two specimens collected by fishers at Gerona (Spain) from a white coral thanatocoenosis, at a depth of 200 – 250 m; those specimens were at first classified as Lepidopleurus alveolus (M. Sars in Lovén, 1846) by Van Belle (1975). This species is little known and scarcely recorded, and the specimens are often difficult to identify, not only because of their small size, but also because of their precarious condition of preservation, as they are mostly found in detritus and therefore coiled up, encrusted, sometimes with a barely visible tegmentum. The fossil record of Leptochiton geronensis is limited to a single tail valve from the Pleistocene of Riparbella (Dell’Angelo & Forli 1995 a). Comparisons. Lepidopleurus geronensis Kaas & Van Belle 1985 is similar to L. xanthus Kaas & Van Belle, 1990, an Atlantic species recently recorded also in the Mediterranean from S. Lucia Bank (Dell’Angelo & Smriglio 1999), a species not treated here because not known as a fossil. The tegmentum of both species presents the same characteristic sculpture of roundish granules well-separated from each other, but L. geronensis differs from L. xanthus by the anterior profile of the intermediate valves (rounded in L. xanthus, semicarinate in L. geronensis), the postmucronal slope (slightly concave in L. xanthus, straight in L. geronensis), and above all by the accessory plate of the second lateral tooth of the radula (monocuspidate in L. xanthus, bicuspidate in L. geronensis) (Dell’Angelo & Smriglio 1999).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD84E280FADFDD468CE916C.taxon	distribution	Distribution: Pleistocene: central Mediterranean, Italy: Riparbella (Dell’Angelo & Forli 1995 a; Dell’Angelo et al. 2001 a). Recent: Mediterranean Sea: Gerona, Spain (Kas & Van Belle 1985 a), Tuscan Archipelago: Formiche di Grosseto (Della Bella & Dell’Angelo 1985), Messina Strait (Scuderi & Dell’Angelo 1997; Dell’Angelo et al. 1998 a), and Ustica Island (Castriota et al. 2005).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD74E290FADFB3369F19321.taxon	description	Fig. 17	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD74E290FADFB3369F19321.taxon	materials_examined	Type material. Holotype: MZB 49985, one intermediate valve (Figs 17 A – C), width 2.08 mm. Paratypes: MZB 49984 (tail valve from Bussana, Fig. 17 H), MSNG 56535 (intermediate valve from Caranchi), BD (tail valve from Bussana, BD 377, Figs 17 E – G), MP (intermediate valve from Rio S. Antonino, Fig. 17 D), MS (tail valve from Rio S. Antonino). Type locality. Genova Sestri (Liguria, Italy). Type stage. Lower Pliocene (Zanclean). Material examined. Lower Pliocene: Italy: type material plus Bussana: 4 valves (BD 379), Genova Sestri: 1 valve (BD 380), Rio S. Antonino: 2 valves (MP), Rio Torsero: 1 valve (BD 381). Maximum width of the valves: -- / 2.1 / 3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD74E290FADFB3369F19321.taxon	description	Description. Valves small. Head valve not available. Intermediate valve broadly rectangular, width greater than 3 times the length, rounded in anterior profile, not much elevated (H / W = 0.24), anterior margin straight, side margins slightly rounded, posterior margin almost straight, apex indistinct, lateral areas scarcely differentiated, with some concentric growth lines. Tail valve elliptical (W / L = 1.66), anterior margin slightly convex, mucro central, prominent, antemucronal slope slightly convex, postmucronal slope concave just underneath mucro. Tegmentum rough, space between striae of granules large, growth lines always evident. LA and PMA sculptured with rather irregular roundish- subquadrangular granules, well separate from each other, arranged in irregular radial series (LA 5 – 6, PA 50 – 52). CA and AMA sculptured with rather irregular roundish- subquadrangular granules, well separate from each other, arranged in irregular longitudinal series (CA ca. 30 – 33, one or two tending to bifurcate, AA ca. 38 – 40, gradually converging posteriorly, some bifurcate), irregular stem structure between granules. Each granule with a central megalaesthete and several micraesthetes irregularly disposed along margin, more or less of same size. Articulamentum without insertion laminae, apophyses wide, triangular in intermediate valves, trapezoidal in tail ones, apical area expanded with straight anterior margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD74E290FADFB3369F19321.taxon	discussion	Remarks. No additional material has been found since the original description. Dell’Angelo et al. (2018 a) attributed tentatively to Leptochiton josei a single tail valve from the lower Miocene of Noaillan (France). Despite the existing similarity with L. josei, it is not possible to identify with certainty this valve, that is discussed and figured in the section dedicated to the “ Species of unclear taxonomic position ”, and it is left in open nomenclature as Leptochiton sp. Comparisons. This species has a certain resemblance to Leptochiton scabridus (Jeffreys, 1880), with which it shares the well separated granules, but which differs from L. josei in having a different and characteristic sculpture, the tegmentum presents a rough surface, on which the granules are extended into a body usually formed by two or three longitudinal varices that become unified or merge together along the external margin of the valve (see above).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD74E290FADFB3369F19321.taxon	distribution	Distribution. Lower Pliocene: central Mediterranean, Italy: Liguria: Bussana, Genova Sestri, Rio S. Antonino, Rio Torsero (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD64E2B0FADF9FC6BEF94EC.taxon	description	(Fig. 18)	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD64E2B0FADF9FC6BEF94EC.taxon	materials_examined	Type material. Holotype, MGPT-PU 108787 (tail valve, width 3.5 mm, Figs 18 A – D). Paratypes from the type locality: MGPT-PU 108788 (head valve, Fig. 18 E); MZB 32033 – 32034 (2 intermediate valves, Figs 18 F – G); NHMW 2014 / 0451 / 0001 – 0002 (intermediate and tail valves); ZISP 2226 – 2227 (2 intermediate valves); MSNG 57980 (1 head and 1 intermediate valves); MZPD MAL 2074 – 2476 (1 head and 2 intermediate valves, Fig. 18 H). Paratype from the Langhian of Moncasale di Casina, “ Fosso di Moncasale ” (Reggio Emilia): MZPD MAL 2077 (intermediate valve). Type locality. Torrente Cinghio, Parma (Emilia-Romagna, Italy). Type stage. Miocene, Tortonian (Termina Formation), a massive dark grey claystone containing few carbonized wood remains that are concentrated within a small lens-shaped feature. Material examined. Miocene (Langhian): Italy: Moncasale di Casina: paratype. Miocene (Tortonian): Italy: Torrente Cinghio: type material plus 100 valves (BD 382, LB). Maximum width of the valves: 2.4 / 3.4 / 3.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD64E2B0FADF9FC6BEF94EC.taxon	description	Description. Head valve semioval, posterior margin widely V-shaped, slope concave. Intermediate valve broadly rectangular (W / L = 2.37), carinate in anterior profile, elevated (H / W = 0.5), anterior and posterior margins almost straight, side margins rounded, apex indistinct, lateral areas scarcely differentiated. Tail valve more than semicircular, anterior margin convex, mucro anterior, not prominent, antemucronal slope convex, postmucronal slope a little concave just underneath mucro. Tegmentum slightly rough, uniformly sculptured with rather irregular roundish, elevated granules (diameter 40 – 50 μm, up to 70 μm in more elongate granules), well separate from each other, irregularly arranged, except apex portion of HV with concentric ridges, granules becoming more irregularly elongate towards JA of intermediate valves, but always well separated, not coalescing. Each granule with a subcentral megalaesthete, and 2 – 4 micraesthetes, up to 5 – 6 in more elongate granules. Articulamentum without insertion laminae, apophyses small, triangular in intermediate valves, somehow trapezoidal in tail ones, widely separated by a large jugal sinus.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD64E2B0FADF9FC6BEF94EC.taxon	discussion	Remarks. The fossil record of Leptochiton lignatilis Dell’Angelo, Bertolaso & Sosso in Bertolaso, Garilli, Parrinello, Sosso & Dell’Angelo, 2015 is limited to the Miocene of N. Italy. No additional material has been found since the original description. In the material studied from Torrente Cinghio there are a few valves complete or sufficiently complete (about a dozen), the others are small fragments (of about 1 – 1.5 mm or less of width), with clearly visible tegmentum granules, but even difficult to identify as head, intermediate or tail valves. The small fragment of the intermediate valve from Moncasale di Casina has a sculpturing matching that of the material from Torrente Cinghio and is compatible with an attribution as Leptochiton. Considering the similarities of the sculpture, this valve was provisionally considered conspecific with the material from Torrente Cinghio. In spite of their considerable presence in recent wood-fall, chitons clearly associated with sunken wood are rare in the fossil records and are only represented by Leptochiton species. A list of these records is reported by Bertolaso et al. (2015), and the description of Leptochiton lignatilis from the Miocene of Italy adds a further record to the scarce number of chitons in fossil wood-fall communities. Comparisons. A comparison of Leptochiton lignatilis with the other Leptochiton specie known from sunken woods (Sigwart & Sirenko 2012) has been made by Bertolaso et al (2015). The closest species is Leptochiton kurnilatus Kaas, 1985 from the Island of Réunion, carinate and elevated (H / W = 0.57), but differs by the different shape and profile of the tail valve, with the posterior mucro at about three quarters of the valve length.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD64E2B0FADF9FC6BEF94EC.taxon	distribution	Distribution. Middle Miocene: Proto-Mediterranean Sea (Langhian): North Italy: Moncasale di Casina (Bertolaso et al. 2015). Upper Miocene: Proto-Mediterranean Sea (Tortonian): North Italy: Torrente Cinghio (Bertolaso et al. 2015).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	description	Fig. 19	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	materials_examined	Type material. Holotype: MSNG 62630, intermediate valve, width 2.4 mm (Figs 19 E – H). Paratype 1: MSNG 62631, head valve, width 1.5 mm (Figs 19 A – B). Paratype 2: MSNG 62632, tail valve, width 1.4 mm (Figs 19 K – L). Type locality. Poggio alla Fame (Tuscany, Italy). Type stage. Pliocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	etymology	Etymology. The name honors Micaela Prudenza (Italy), for her contribution to the study and research of fossils from the Pliocene of Liguria and Tuscany.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	materials_examined	Material examined. Pliocene: Italy: Tuscany: Poggio alla Fame: type material plus 3 valves (BD 383); Sicily: Trappeto: 1 valve (BD 384). Pleistocene: Italy: Tuscany: Riparbella: 2 valves (BD 385); Puglia: Gallipoli: 1 valve (PC); Calabria: Pecoraro: 1 valve (BD 386); Sicily: Selinunte, Casa Parrino: 4 valves (BD 387, Figs 19 I – J). Greece: Kyllini: 5 valves (BD 388, Figs 19 C – D, DGUP). Maximum width of the valves: 1.5 / 2.4 / 1.4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, rounded, moderately elevated, apex inconspicuous, tail valve semicircular, mucro subcentral, not prominent. Tegmentum rough, sculptured with well raised, distinctly separated granules, elliptical and irregularly arranged in HV, LA, PMA, fungiform with 2 – 3 varices, arranged in longitudinal series in CA, AMA, each granule with 4 – 6 aesthetes irregularly disposed in CA and AMA, all same size.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, slope straight. Intermediate valves broadly rectangular, elongated (W / L = 2.36 – 3.40), rounded in anterior profile, moderately elevated (H / W = 0.33 – 0.37), anterior margin stright, side margins rounded, posterior margin straight, apex inconspicuous, lateral areas hardly or not raised. Tail valve semicircular, anterior margin almost straight or slightly convex, mucro not prominent, in subcentral position, antemucronal slope slightly convex, postmucronal slope slightly concave. Tegmentum rough. HV, LA and PMA sculptured with well raised, distinctly separated elliptical granules, irregularly arranged, close set and forming concentric lines. CA and AMA sculptured with distinctly separated fungiform granules, with a roundish body (up to 38 µm) extended with 2 – 3 longitudinal varices, more widely spaced and arranged in longitudinal series (CA 50 – 55, AMA 40 – 43), less parallel and slightly divergent in AMA, with a regular quincuncial pattern displayed by granules of neighboring rows. Each granule with 4 – 6 aesthetes irregularly disposed in CA and AMA, all aesthetes of same size. Articulamentum without insertion laminae, with apophyses small, sharply triangular, widely separated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	discussion	Remarks. Much of the fossil material studied had already been reported in previous publications, and attributed to Leptochiton cimicoides (Monterosato, 1879), the least known of the four species described by Monterosato in the Mediterranean Sea. However, some differences compared to living specimens of L. cimicoides had already been highlighted, e. g., for the material from Kyllini’s Pleistocene [Garilli et al. 2005: 130 “ The valves found agree well with the characteristics of the species, except for the number of longitudinal series of granules in central areas of intermediate valves, about 50 (in our valves) vs. 30 (as reported by Dell’Angelo & Smriglio, 1999) ”], but interpreted as intraspecific variation by the authors. The recent discovery of abundant material from the Pliocene of Poggio alla Fame (Tuscany, Italy) made it possible to better evaluate these differences and to attribute all the fossil material to a new species; in our opinion, L. cimicoides is at present only known as a living species in the Mediterranean Sea and is not included in the present study. The fossil record of Leptochiton prudenzae sp. nov. refers to the Pliocene (Poggio alla Fame, Trappeto) and Pleistocene of Italy (Gallipoli, Pecoraro, Riparbella, Selinunte) and Greece (Kyllini, Rhodes). Comparisons. The closest species is Leptochiton cimicoides (Monterosato, 1879), from which L. prudenzae sp. nov. differs mainly by the different number of longitudinal series of granules in central and antemucronal areas (50 – 55 vs. 30), the space between the striae of granules narrower, the subcentral mucro in the tail valve (in posterior position in L. cimicoides).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD44E2C0FADFDB06E4393E1.taxon	distribution	Distribution. Pliocene: central Mediterranean, Italy: Poggio alla Fame (this study), Trappeto (Dell’Angelo & Palazzi 1989). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece: Rhodes (Koskeridou et al. 2009). Pleistocene: central Mediterranean, Italy: Gallipoli, Pecoraro, Riparbella, Selinunte (Dell’Angelo & Palazzi 1989; Dell’Angelo & Forli 1995 a; this study), Greece: Kyllini (Garilli et al. 2005).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD34E2D0FADF8BF6F3D9268.taxon	description	Fig. 20	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD34E2D0FADF8BF6F3D9268.taxon	materials_examined	Type material. Holotype: MZB 31028 (1 intermediate valve, Figs 20 B – F). Paratypes: MZB 31029: 2 valves (1 head and 1 tail, Figs 20 A, 20 G – H); BD 4662: 4 valves (2 intermediate and 2 tail). Type locality. Salice, Messina (Sicily, Italy). Type stage. Lower Pleistocene. Material examined. Lower Pleistocene: Italy: Sicily: Salice: type material plus 10 valves (BD 389). Maximum width of the valves: 4.3 – 4.9 – 3.9 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD34E2D0FADF8BF6F3D9268.taxon	description	Description. Head valve semicircular. Intermediate valves broadly rectangular (W / L = 1.66), rounded in anterior profile, elevated (H / W = 0.51), anterior margin slightly convex, side margins rounded, posterior margin straight, apex inconspicuous, lateral areas not raised. Tail valve semicircular, anterior margin almost straight, mucro subcentral, not prominent, antemucronal slope slightly convex, postmucronal slope slightly concave. Tegmentum rough, concentric lines of growth well visible, uniformly sculptured with well raised, neatly separated roundish / polygonal granules, randomly arranged, diameter 70 – 90 μm, aesthetes not clearly visible, a central megalaesthete and 2 – 3 micraesthetes (but there should be more) around the border of some granules. Articulamentum without insertion laminae, apophyses small, sharply triangular, widely separated by a large jugal sinus, apical area expanded, with bisinuate anterior margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD34E2D0FADF8BF6F3D9268.taxon	discussion	Remarks. The fossil record of Leptochiton salicensis (Dell’Angelo & Bonfitto, 2005) is limited to the Lower Pleistocene of Salice. Dell’Angelo et al. (2015) attributed tentatively to Leptochiton salicensis two valves, a head from the lower Miocene of Rocco di Passerano (Italy) and a tail from the Miocene (Tortonian of Borelli (Italy). Despite the existing similarity with L. salicensis, it is not possible to identify with certainty these valves, that are discussed and figured in the section dedicated to the “ Species of unclear taxonomic position ”, and are left in open nomenclature as Leptochiton sp. Comparisons. See Tab. 5 for a comparison with the Leptochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD34E2D0FADF8BF6F3D9268.taxon	distribution	Distribution. Lower Pleistocene: central Mediterranean: Italy: Salice (Dell’Angelo & Bonfitto 2005).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD14E2F0FADFF056A9495C4.taxon	description	Fig. 21	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD14E2F0FADFF056A9495C4.taxon	materials_examined	Type material. Holotype KNVSM, specimen 7.7 x 3.9 mm. Paratypes: KNVSM (2 paratypes); ZMO, type collection D 33690 (1 paratype); K 4914 (1 paratype). Type locality. Asfjorden, Trondheimsfjord (Norway). Material examined. Pleistocene: Italy: Calabria: Archi: 4 valves (BD 390, Figs 21 B – D), Bovetto: 1 valve (BD 391), Cannitello: 1 valve (FG), San Procopio: 3 valves (BD 392, Figs 21 G – H), Venetico Marina: 1 valve (BD 393, Fig. 21 A); Sicily: Pezzo: 1 valve (BD 394), Salice: 17 valves (BD 395, Figs 21 E – F). Recent: Italy: Tuscan Archipelago, S. Lucia Bank: 2 specimens (BD 396, Figs 21 I – L). Maximum width of the valves: - / 4 / 3.4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD14E2F0FADFF056A9495C4.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 1.90 – 2.12), rounded in anterior profile, elevated (H / W = 0.46 – 0.55), anterior margin slightly convex, side margins rounded, posterior margin almost straight, apex not evident, lateral areas slightly raised with concentric lines of growth. Tail valve almost semicircular (W / L = 1.84 – 1,90), anterior margin almost straight or slightly convex, mucro not prominent, subcentral or a little anterior, antemucronal slope convex, postmucronal slope concave directly behind the mucro. Tegmentum rough, space between striae of granules large. HV, LA, PMA sculptured with small elliptical granules randomly arranged, with 2 – 3 stems in apical part. CA and AMA sculptured with more roundish and more compact granules arranged in longitudinal series (CA 50 – 60, AMA ca 45), with a regular quincuncial pattern displayed by granules of neighbouring rows. Each granule with a variable number of aesthetes, normally three lined up along diameter, a central megalaesthete and two micraesthetes at extremities in HV, LA and PMA, and six with a central megalaesthete and two or more micraesthetes on two sides in CA and AMA. Articulamentum without insertion laminae, weakly developed, apophyses small, triangular, wide apart in intermediate valves, trapezoid in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD14E2F0FADFF056A9495C4.taxon	discussion	Remarks. Leptochiton sarsi Kaas, 1981 was described by Kaas (1981) in his revision of Leptochiton living along the Scandinavian coasts. By studying a lot of specimens, identified as L. cancellatus (Sowerby, 1840), Kaas found that specimens described and figured by G. O. Sars (1878: 111, pl. 7, figs 6 a – 6 h) as L. cancellatus, did not agree with the species described by Sowerby (1840 a: figs 104, 104 a – 104 b, 105; 1840 b: 4), and these were described as the new species L. sarsi Kaas, 1981, along with an elaboration of the differences between the two taxa (Kaas 1981: table 2). The distribution of L. cancellatus was restricted to the British Isles, the Atlantic coasts of France, Spain and Portugal, and the Mediterranean Sea, while L. sarsi appeared to be continuously distributed along the Scandinavian coast, from Bohuslän (Sweden) up to Finmark (Norway) at a depth of 40 to over 700 m. Leptochiton sarsi has been recently found in the Mediterranean Sea, on the basis of two specimens from S. Lucia Bank, 500 m depth (Dell’Angelo et al. 2009). The species was already previously reported as loose valves, considered subfossil (Giovine & Dell’Angelo 1993: an intermediate valve from biogenic sands at Cannitello, Reggio Calabria, at 150 m depth. The citation of Dell’Angelo & Giusti (2000: many valves from the Southern Ligurian Sea, between Capraia Island. and Capo Corso, 350 / 500 m depth) should be referred Leptochiton corticellii sp. nov.). Moreover, the species was already reported as a fossil in bathyal Pleistocene outcrops of S. Italy: Archi (figured by Dell’Angelo & Palazzi 1989: pl. 21, figs 1 – 2), San Procopio (figured by Dell’Angelo & Palazzi 1989: pl. 25, figs 1 – 3), Salice (Dell’Angelo & Palazzi 1989), and Venetico Marina (Palazzi & Villari 1994). Comparisons. Leptochiton sarsi differs from L. cancellatus (Sowerby, 1840) mainly by the different sculpture, the more spaced rows of granules on CA and AMA, the granules larger and not connected. Leptochiton sarsi is also closest to Leptochiton corticellii sp. nov.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD14E2F0FADFF056A9495C4.taxon	distribution	Distribution. Pleistocene: central Mediterranean, S. Italy: Archi, Bovetto, Cannitello, Pezzo, San Procopio, Salice, Venetico Marina (Dell’Angelo & Palazzi 1989; Giovine & Dell’Angelo 1993; Palazzi & Villari 1994; this study). Recent: N. Atlantic Ocean, along the Scandinavian coast from Sweden to Norway (Hansson 1998; Kaas 1981; Kaas & Van Belle 1985 a), and Mediterranean Sea: Tuscan Archipelago: S. Lucia Bank (Dell’Angelo et al. 2009).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD04E510FADFAC76EDA9108.taxon	description	Fig. 22	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD04E510FADFAC76EDA9108.taxon	materials_examined	Type material. Syntypes: Goodrington, Torbay: 1 specimen (USNM 177391); Jersey: 15 specimens (USNM 177392) (fide Warén 1980). Type locality. Goodrington, Torbay, England and Jersey (Channel Islands). Material examined. Upper Miocene: Italy: Montegibbio: 1 valve (MZB 32013), Rio di Bocca d’Asino: 3 valves (BD 397, MZB 32012). Lower Pliocene: Italy: Liguria: Genova Sestri: 1 valve (MZB 45702, Fig. 22 D). Pliocene: Portogallo: 14 valves (BD 238, GeoFCUL VFX. 03.333, GeoFCUL VFX. 03.350, RGM. 1363999 – 1364000, MNHN. F. A 81982, Figs 22 G – L). Italy: Tuscany: Poggio alla Fame: 2 valves (BD 258). Pleistocene: Italy: Calabria: Pecoraro: 2 valves (BD 398, Figs 22 E – F). Greece: Kyllini: 2 valves (BD 399, DGUP, Figs 22 A – C). Maximum width of the valves: 1.4 / 3.7 / 2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD04E510FADFAC76EDA9108.taxon	description	Description. Valves small, fragile. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular, length more or less three times the width (W / L = 2.88 – 3.30), rounded in anterior profile, moderately elevated (H / W = 0.25 – 0.32), anterior margin straight, side margins rounded, posterior margin straight, apex not indicated, lateral areas hardly raised but clearly indicated. Tail valve semicircular (W / L = 1.77 – 1.90), front margin straight to slightly convex, mucro about central, somewhat swollen, antemucronal slope convex, postmucronal slope concave. Tegmentum rough, space between striae of granules large, some growth lines, more evident in LA. HV, LA and PMA sculptured with irregular subquadrangular / subrhomboidal granules well separated from each other, arranged in radiating series (HV 36 – 48, LA 7 – 8, PMA 30 – 36). CA and AMA sculptured with irregular subquadrangular / subrhomboidal granules well separated from each other, arranged in longitudinal series (CA 24 – 30), tending to become slightly converging near outer sides and looking like square mesh, granules extended with 2 – 3 longitudinal varices that become unified or merge together along external margin of valve. Each granule with a central megalaesthete and some micraesthetes irregularly disposed. Articulamentum lacking insertion laminae, apophyses small, sharply triangular, widely separated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD04E510FADFAC76EDA9108.taxon	discussion	Remarks. Leptochiton scabridus (Jeffreys, 1880) was described but not figured by Jeffreys and is quoted by Pilsbry (1894) only in Appendix II (“ Insufficiently described Chitons, and species of unknown generic position ”), which, however, reports the original description. It was first figured by Sykes (1894), while Thiele (1909) described the radula and the girdle on a topotypic specimen from Jersey. The species was described with a distribution limited to a small zone of the British Channel along the English and French coasts, and subsequently it was rarely found in other areas of the Mediterranean, initially confused with L. cancellatus (Dell’Angelo & Palazzi 1989; Dell’Angelo & Smriglio 1999). Possibly because of uncorrect attributions to the close species Leptochiton cancellatus (see Dell’Angelo & Palazzi, 1989: 61), historical records are doubtful and require confirmation. The fossil record of Leptochiton scabridus is limited thus far to the upper Miocene of Italy (Dell’Angelo et al. 2015 a), the Pliocene of Italy (Dell’Angelo et al. 2013) and Portugal (Dell’Angelo et al. 2022), and the Pleistocene of Italy (this study) and Greece (Garilli et al. 2005). Comparisons. The species that have the greatest similarities with Leptochiton scabridus are those that share the sculpture of the tegmentum formed by striae of granules well separated from each other (L. serenae Dell’Angelo, Piccioli Resta & Bonfitto, 2007 and L. josei Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013). These species differ from each other for a series of characteristics relating to the shape and sculpture of the valves, well highlighted in Tab. 2.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFD04E510FADFAC76EDA9108.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Montegibbio, Rio di Bocca d’Asino (Dell’Angelo & Palazzi 1989; Dell’Angelo et al. 2015 a). Lower Pliocene: central Mediterranean, Italy: Liguria: Sestri Ponente (Dell’Angelo et al. 2013). Pliocene: northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo et al. 2022); central Mediterranean, Italy: « La Tagliata », Modena (Laghi 1977, as L. cancellatus; Dell’Angelo & Palazzi 1989), Poggio alla Fame (this study). Pleistocene: central Mediterranean, S. Italy: Pecoraro (this study), Greece: Kyllini (Garilli et al. 2005). Recent: Atlantic Ocean: S. W. U. K. and the British Channell (Light & Baxter 1990), France: Bretagne (Van Belle 1972), northern Spain (Urgorri et al. 2017), Portugal (Consolado Macedo et al. 1999), Canary Islands (Hernández & Rolán 2011), Cape Verde Islands (Kaas 1991) and Angola (Dell’Angelo & Smriglio 1999). Mediterranean Sea: Italy: Tuscan Archipelago (Dell’Angelo & Palazzi, 1986), Taranto Gulf and Otranto coast (Dell’Angelo & Palazzi 1986; Baschieri 1994), and many other localities (Dell’Angelo & Palazzi 1986; Dell’Angelo & Smriglio 1999); Greece and Aegean Sea Islands (Strack 1988; Koukouras & Karachle 2005); Malta (Mifsud et al. 1990); Turkey (Ozturk et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAE4E520FADFB146BA1901E.taxon	description	Fig. 23	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAE4E520FADFB146BA1901E.taxon	materials_examined	Type material. Holotype: MZB 23749 (1 tail valve, Isola del Campo, Figs 23 A – D). Paratypes: MZB 23750 (1 intermediate valve, Cutrofiano, Fig. 23 H); BD 4735 (1 intermediate, and 1 tail valve, Isola del Campo, Figs 23 E – G); GPR (1 intermediate valve, Isola del Campo, and 1 tail valve, Cutrofiano). Type locality. Isola del Campo, Gallipoli (Puglia, Italy). Type stage. Lower Pleistocene (Santernian-Aemilian limit). Material examined. Lower Pleistocene: Italy: type material. Maximum width of the valves: -- / 4.7 / 4.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAE4E520FADFB146BA1901E.taxon	description	Description. Head valve not available. Intermediate valves broadly rectangular, subcarinate in anterior profile, moderately elevated (estimated H / W = 0.42), anterior and posterior margins quite straight, side margins slightly rounded, jugal sinus crenulated, apex inconspicuous, lateral areas hardly or not raised, but clearly recognizable by different sculpture. Tail valve semicircular, width more than twice length (W / L = 2.00 – 2.08), anterior margin almost straight, anterior, prominent mucro, backward directed, antemucronal slope convex, postmucronal slope slightly concave. Tegmentum rough, space between striae of granules reduced, growth lines thin near margins of LA and stronger on PMA. LA and PMA sculptured with oval granules, separated from each other, developing obliquely from tegmentum, arranged in radiating series (LA 7 – 8, PMA 36 – 50). CA and AMA sculptured with oval granules, separated from each other, developing obliquely from tegmentum, arranged in longitudinal series (CA 60, AMA 36 – 60). Each granule with 3 – 5 aesthetes of equal width, not differentiated in megalaesthetes and micraestehetes, and not placed in a peculiar order. Articulamentum without insertion laminae, apophyses wide, triangular in intermediate valves, trapezoidal in tail valve, apical area expanded, with straight anterior edge.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAE4E520FADFB146BA1901E.taxon	discussion	Remarks. The species is known only for the type material, and is characterized by its sculpture of granules developing obliquely from the tegmentum, which differs from that of other known living and fossil Mediterranean and Atlantic Leptochiton species. None of the three intermediate valves is complete, so it has been possible only to estimate in one valve the height / width ratio. Comparisons. See Tab. 2 for a comparison with the Leptochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAE4E520FADFB146BA1901E.taxon	distribution	Distribution. Lower Pleistocene: central Mediterranean, S. Italy: Cutrofiano, Isola del Campo (Dell’Angelo et al. 2007 b; Brunetti 2011).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAD4E530FADFAE66B62935E.taxon	description	Fig. 24	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAD4E530FADFAE66B62935E.taxon	materials_examined	Type material. Holotype: MZB 25049 (1 intermediate valve, Figs 24 A – C). Paratypes: MZB 25050 (1 intermediate and 1 tail valve); MME (1 intermediate and 1 tail valve); IRScN IST 6449 (1 intermediate and 1 tail valve); BD 4566 (2 intermediate valves, Fig. 24 I); RM (2 intermediate valves). Type locality. Estepona, Velerín Carretera (Spain). Type stage. Pliocene, lower Piacenzian. Material examined. Pliocene: Spain: Estepona: type material plus 38 valves (BD 400, Figs 24 D – H, 24 J – L). Maximum width of the valves: 3.7 / 4.6 / 3.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAD4E530FADFAE66B62935E.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, slope straight. Intermediate valves broadly rectangular (W / L 2.27 – 2.54), rounded in anterior profile, elevated (H / W = 0.39 – 0.55), anterior margin slightly convex, side margins rounded, posterior margin straight, apex inconspicuous, lateral areas slightly or not raised. Tail valve more than semicircular (W / L = 1.57 – 1.83), anterior margin almost straight, mucro subcentral, prominent, antemucronal slope straight, postmucronal slope concave. Tegmentum rough, uniformly sculptured with well raised, neatly separated, fungiform section granules (width 54 – 65 μm), arranged in bee’s nest structure, i. e. six granules at corners of hexagon and one in the center, suggesting longitudinal or radial, mutually staggered striae, or a trellis-work structure, of diagonal, intersect striae. Each granule with generally 5 aesthetes of equal width, one of which lies in axial plane of prominence with the other four situated two by two at its sides. Articulamentum without insertion laminae, apophyses small, sharply triangular, widely separated by large jugal sinus, apical area expanded, with bisinuate superior margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAD4E530FADFAE66B62935E.taxon	discussion	Remarks. The fossil record of Leptochiton tavianii (Dell’Angelo, Landau & Marquet, 2004) is limited to the Pliocene of Estepona (Spain). The head valve was not known in the material described by Dell’Angelo et al. (2004), and it is here figured (Figs 24 D – E). Comparisons. Leptochiton tavianii is well characterized by the sculpture of tegmentum which differs from closer species in the L. tavianii group spp. by the granules with fungiform section arranged in beehive structure.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAD4E530FADFAE66B62935E.taxon	distribution	Distribution. Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004; this paper).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAB4E550FADFF056F0C9352.taxon	description	Fig. 25	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAB4E550FADFF056F0C9352.taxon	description	p. 354, 412, 417 (fide Malatesta 1962). Lepidopleurus asellus;? Antevs 1928, p. 554 – 555, 646, 678 – 689; Marquet 1984, p. 336, pl. 1, fig. 1. Lepidopleurus cf. asellus; Bellomo & Sabelli 1995, p. 201. Lepidopleurus (Leptochiton) asellus; Malatesta 1962, p. 150, figs 5 – 6; Dell’Angelo & Palazzi 1989, p. 56, pl. 5; Dell’Angelo & Smriglio 1999, p. 43, pls 8 – 9, figs 16 – 17; Dell’Angelo & Giusti 2000, p. 53, figs 1 – 4; Marquet 2002, p. 11, pl. 1, fig. 1. Leptochiton asellus; Kaas 1981, p. 217, fig. 1 – 3; Strack 2010, p. 61, figs 46 – 49; Taviani et al. 2023, p. 3, fig. 2. Leptochiton (L.) asellus; Kaas & Van Belle 1985 a, p. 39, fig. 15; Sturrock & Baxter 1993, p. 49, pls 1 – 6.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAB4E550FADFF056F0C9352.taxon	materials_examined	Type material: Holotype at the Zoological Museum of the University of Copenhagen, a specimen attached to a shell of Modiolus (Kaas & Knudsen 1992: fig. 26). Type locality: Telemark, Kragerö (Norway). Material examined. Lower Pliocene: Belgium: Kallo: 3 valves (BD 401, Figs 25 G – J). Pleistocene, presumably last glacial: Italy: Capraia Island-Capo Corso - 350 / 500 m: 14 valves (BD 402, Figs 25 A – F); off Bari, cruise SE 06: SE 06 - 10: 1 valve; SE 06 - 13: 1 valve; SE 06 - 18: 2 valves; SE 06 - 19: 1 valve; SE 06 - 22: 1 valve; SE 06 - 24: 1 valve; SE 06 - 25: 2 valves; SE 06 - 35: 5 valves; SE 06 - 40: 1 valve; SE 06 - 48: 1 valve; SE 06 - 50: 25 valves (Figs 25 K – L). Maximum width of the valves: 3.6 / 5.2 / 4.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAB4E550FADFF056F0C9352.taxon	description	Description: Head valve almost semicircular. Intermediate valves broadly rectangular, length more than three times the width (W / L = 3.03 – 3.30), moderately elevated (H / W = 0.36 – 0.38), semicarinate in anterior profile, anterior margin straight, lateral margins slightly rounded, posterior margin slightly concave at both sides of not very pronounced but well visible apex, lateral areas little raised. Tail valve semicircular, width about twice length (W / L = 1.88 – 2.05), mucro slightly anterior, postmucronal slope almost straight or slightly concave. Tegmentum rough, space between striae of granules reduced, growth lines often present in variable number. HV, LA and PMA sculptured with minute, low, roundish to oval granules arranged in radial series (HV 70 – 80, LA 15 – 20). CA and AMA sculptured with roundish, united granules arranged in longitudinal striae (CA 70 – 80), displaying a regular quincuncial pattern by granules of neighboring rows, tending to bend slightly towards outer margins. Each granule with aesthetes of same size, one central and 2 – 4 irregularly arranged. Articulamentum lacking insertion laminae, apophyses small, well separate, triangular but tending to become trapezoidal in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAB4E550FADFF056F0C9352.taxon	discussion	Remarks: Chiton asellus Gmelin, 1791 was originally described and inadequately illustrated by Chemnitz (1785) based on a Norwegian specimen attached to a shell of Modiolus, “ ex Museo Spengleriano ”. This description is not valid, Chemnitz’s work not being strictly binomial. The taxon was validated by Gmelin (1791), who added nothing new to the description, and was later better defined by Spengler (1797). This species has a complicated taxonomic history, and a thorough discussion of the interpretations given by various authors was provided by Kaas (1981) and Kaas & Knudsen (1992). The fossil records of this species are rather scarce and must be accepted with caution; historical records are doubtful and require confirmation. The intermediate valves reported from the lower Pliocene of Kallo (Belgium) by (Marquet 1984, 2002) show a more rounded anterior profile (Fig. 25 H), not quite semicarinate as they should be (Fig. 25 D), but all the other features of the plates and of the sculpture agree with those of Leptochiton asellus, so we confirm this attribution. Many valves of Leptochiton asellus (Gmelin, 1791) from Sea-bottom sampling at bathyal depths off the Apulian margin (southwestern Adriatic Sea, SE 06 cruise) have recently been studied by Taviani et al. (2023), and this finding is quite significant, extending the presence of this species in the Italian Pleistocene, hitherto known only for a few finds. Comparisons. See Tab. 3 for a comparison with the Leptochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAB4E550FADFF056F0C9352.taxon	distribution	Distribution. Lower Pliocene: North Europe, Belgium: Kallo (Marquet 1984, 2002); NE Atlantic: U. K. (Wood 1842, 1848; Reid 1890; Malatesta 1962); Pleistocene: North Atlantic: Netherlands (Strack 2010); central Mediterranean, Italy: Pezzo (Bellomo & Sabelli 1995). Pleistocene, presumably last glacial: North Atlantic: Sweden and Norway (Brogger 1901; Antevs 1917, 1928); central Mediterranean, Italy: Capraia Island-Capo Corso - 350 / 500 m (Dell’Angelo & Giusti 2000), cruise SE 06, off Bari (Taviani et al. 2023). Recent: North Europe: from Spitsbergen and the Barents Sea South along the Scandinavian coasts (Dons 1934; Hansson 1998); lceland (Sneli & Gudmundsson 2018) (and perhaps Greenland); NE Atlantic Ocean: all around the British Islands and Ireland (Light & Baxter 1990; McKay & Smith 1979), coasts of France (Van Belle 1972; Kaas 1979), S. to Spain (Borja 1987; Rolan Mosquera et al. 1990; Urgorri et al. 2017) and Portugal (Consolado Macedo et al. 1999) (Kaas 1981; Kaas & Van Belle 1985 a). Mediterranean Sea: a single record from the Limnos Island, Greece (Mifsud & Ovalis 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAA4E570FADF953685E975C.taxon	description	Fig. 26	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAA4E570FADF953685E975C.taxon	materials_examined	Type material. Holotype MNHN. F. A 67071, intermediate valve, width 6.3 mm (Figs 26 B – D). Paratypes: MNHN. F. A 67072 – A 67074 (head, intermediate and tail valves, Figs 26 A, 26 G – H); NHMW 2017 / 0108 / 0008 – 0010 (head, intermediate and tail valves, respectively); RGM. 1008365, intermediate valve (Figs 26 E – F). Type locality. Saint-Clément-de-la-Place, Anjou, France. Type stage. Miocene (Tortonian). Material examined. Upper Miocene (Tortonian): Saint-Clément-de-la-Place: type material plus 36 valves (MNHN. F. A 67075, NHMW 2017 / 0108 / 0011, RGM. 1008366, RGM. 1008398, BD 135). Maximum width of the valves: 6.3 / 7 / 4.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAA4E570FADF953685E975C.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, notched in middle, slope straight. Intermediate valve broadly rectangular, width more than three times the length (W / L = 3.32 – 3.40), rounded to subcarinate in anterior profile, moderately elevated (H / W = 0.22 – 0.39), anterior margin slightly convex, side margins rounded, posterior margin straight, apex not developed, lateral areas slightly raised. Tail valve almost semicircular, width ca. two times the length (W / L = 1.96), anterior margin convex, mucro in anterior position, antemucronal slope slightly convex, postmucronal slope concave, more pronounced concavity just behind mucro, antemucronal area short, less than one – third of length of postmucronal area. Tegmentum rough. HV, LA and PMA sculptured with radial granular striae intersected by concentric growth lines, with some elevated pustules irregularly distributed on striae (HV ca. 76, LA ca. 12, PMA ca. 65), concentric growth lines continuing across central area, where they weaken, becoming inconspicuous. CA and AMA sculptured with longitudinal granular striae, weakening anteriorly, not reaching anterior margin, subobsolete in jugal area, which appears smooth without magnification. Each granule with a central megalaesthete and 5 – 6 micraesthetes irregularly disposed. Articulamentum without insertion laminae, apophyses well developed, bluntly triangular with a rounded margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAA4E570FADF953685E975C.taxon	discussion	Remarks. The record of Leptochiton lateropustulosus Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is to date limited to the original material from the Miocene (Tortonian) of France (Dell’Angelo et al. 2018 b). The intermediate valves are highly variable in shape (H / W = 0.22 – 0.39, 0.34 in holotype) and most markedly in the sculpture of PA and JA, where many granular longitudinal striae do not reach the anterior margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFAA4E570FADF953685E975C.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA84E580FADFD2068EA9420.taxon	description	Fig. 27	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA84E580FADFD2068EA9420.taxon	materials_examined	Type material. Holotype: MNHN. F. A 67069, intermediate valve, width 2.5 mm, Fig. 27 A – D. Paratypes: MNHN. F. A 67070, intermediate valve; RGM. 1310174, intermediate valve from Beugnon. Type locality. Saint-Clément-de-la-Place (Anjou, France). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): France: Saint-Clément-de-la-Place: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA84E580FADFD2068EA9420.taxon	description	Description. Head and tail valves unknown. Intermediate valve broadly rectangular, very elongated (W / L = 3.48), rounded to subcarinate in anterior profile, moderately elevated (H / W = 0.29), anterior and posterior margins almost straight, side margins slightly rounded, apex not developed, lateral areas scarcely differentiated. Tegmentum rough, space between striae of granules narrow. LA sculptured with seven irregular radial series of widely spaced roundish granules. CA sculptured with 30 longitudinal series of widely spaced roundish granules, of ca. 50 µm of diameter. Each granule with a central megalaesthete and 5 – 8 micraesthetes irregularly disposed along margin. Articulamentum without insertion laminae, apophyses triangular.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA84E580FADFD2068EA9420.taxon	discussion	Remarks. To date, Leptochiton parvus Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is only known upon three small intermediate valves from Saint-Clément-de-la-Place and one from Beugnon, Miocene (Tortonian) of France (Dell’Angelo et al. 2018 b) Comparisons. Leptochiton parvus is closest to the other species of the Leptochiton cancellatus group, from which it differs by the tegmentum sculpture with a greater number of granular longitudinal striae and smaller interspaces.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA84E580FADFD2068EA9420.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place, Beugnon (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA74E590FADFEFC6ED89490.taxon	description	Fig. 28	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA74E590FADFEFC6ED89490.taxon	materials_examined	Type material. Holotype MNHN. F. A 67076, tail valve, width 3 mm (Figs 28 A – E). Paratype: MNHN. F. A 67077, intermediate valve, width 3 mm (Figs 28 F – H). Type locality. Renauleau (Anjou, France). Type stage. Miocene (Tortonian). Material examined. Upper Miocene (Tortonian): France: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA74E590FADFEFC6ED89490.taxon	description	Description. Head valve unknown. Intermediate valve broadly rectangular, very elongated (W / L = 3.09), subcarinate in anterior profile, moderately elevated (H / W = 0.33), posterior margin straight, apices not indicated, lateral areas not raised, delimited only by change in sculpture compared to central area. Tail valve oval, mucro not protruding in anterior position, antemucronal slope almost straight, postmucronal slope slightly concave. Tegmentum rough, granules well separated from each other, growth lines not very evident. LA and PMA sculptured with granules randomly arranged. CA and AMA sculptured with longitudinal series of widely spaced roundish and elevated granules (CA 50 – 55, AMA 40, more irregular), converging towards outer margins. Each granule with a central megalaesthete and 5 – 6 micraesthetes irregularly disposed along margin. Articulamentum without insertion laminae, apophyses wide, not well preserved.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA74E590FADFEFC6ED89490.taxon	discussion	Remarks. The fossil record of Leptochiton renauleauensis Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is limited to the Miocene (Tortonian) of France (Dell’Angelo et al. 2018 b). The two valves found are not complete, quite worn and some characters are not observable. The longitudinal series of granules on the central area of intermediate valves and the antemucronal area of tail valve are quite irregular, so much to give the impression of being randomly arranged in certain areas. Despite this, the sculpture is different from that of the other Leptochiton species known from the European Neogene, and characteristic enough to have been described as a new species by Dell’Angelo et al. (2018 b). We estend the stratigraphic distribution of Leptochiton renauleauensis to the lower Pliocene, attributing to this species also the valves from Kallo (Belgium) illustrated by Marquet (2002) as Lepidopleurus (Leptochiton) sp. Comparisons. The characteristics of the sculpture of Leptochiton renauleauensis agree with the belonging of the species to the Leptochiton cimicoides group.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA74E590FADFEFC6ED89490.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Renauleau (Dell’Angelo et al. 2018 b). Lower Pliocene: North Europe, Belgium: Kallo (Marquet 2002).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA64E5A0FADFC8C6F1B931E.taxon	description	Fig. 29	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA64E5A0FADFC8C6F1B931E.taxon	description	Dell’Angelo & Smriglio 1999, p. 72. Leptochiton srameki; Macioszczyk 1988, p. 51, pl. 1 fig. 9; Dell’Angelo et al. 2018 b, p. 52, tab. 17.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA64E5A0FADFC8C6F1B931E.taxon	materials_examined	Type material. Holotype SMF 1413 a, tail valve, width 3.2 mm (Fig. 29 A), Paratype SMF 1413 b, head valve, width 3.8 mm (Figs 29 B – C), Paratype, SMF 1413 c _ 2, intermediate valve, width 4 mm (Fig. 29 D), Paratype, SMF 1413 c _ 1, intermediate valve, width 3.4 mm (Figs 29 E – F). Type locality. Kostej (Romania). Type stage. Middle Miocene. Material examined. No actual material available, only descriptions and illustrations from the literature.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA64E5A0FADFC8C6F1B931E.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, notched in middle. Intermediate valves broadly rectangular, carinate in anterior profile, moderately elevated, anterior margin slightly convex, posterior margin almost straight, apex not developed, lateral areas slightly raised. Tail valve semicircular, mucro in central position. Tegmentum rough. HV and PMA sculptured with radial striae of oval granules (HV 50, PA 35), not all starting from apex, but tending to branch, LA with oval granules seeming arranged in concentric growth lines and, in addition, in radial striae. CA and AMA sculptured with longitudinal striae of closely spaced roundish granules (CA 20 on each side, striae more closely spaced in jugum than on slopes, AMA 22 – 26, converging towards outer margins and quite far from each other). Articulamentum without insertion laminae, with small apophyses, widely separated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA64E5A0FADFC8C6F1B931E.taxon	discussion	Remarks. Šulc (1934) described two species, L. srameki and L. boettgeri, considered cospecific by Dell’Angelo & Palazzi (1989), that considered L. srameki as a junior synonym of L. boettgeri. Šulc himself considered the two species very similar, highlighting as difference that “ not all the radial chains of granules start from the apex ”, but tend to branching (in L. boettgeri). Leptochiton boettgeri was established by Šulc (1934) on the basis of four valves (Figs 29 A – F) from the Boettger collection, previously determined by O. Boettger as Chiton sp., while L. srameki was determined by Šulc (1934) on the basis of a single tail valve (Fig. 29 H). Subsequently Bałuk (1971, 1984) attributed to L. srameki two intermediate valves from Korytnica (Poland) (Fig. 29 G), remarking that an exhaustive comparison is unfeasible. Another intermediate valve from Węglinek (Poland) was reported by Macioszczyk (1988), a valve consistent with the material described by Bałuk (1971, 1984). Dell’Angelo & Palazzi (1989) recorded the strong affinities existing between the two fossil species described by Šulc (1934) and the recent Mediterranean species Leptochiton bedullii Dell’Angelo & Palazzi, 1986, and put L. bedullii in synonymy with L. boettgeri, 1934, on the basis of a direct comparison between a tail valve (paratype) of L. bedullii and the holotype of L. boettgeri. Kaas & Van Belle (1994) consider the two species distinct, above all on the basis of differences between the Miocene molluscan faunas and the Recent. Dell’Angelo & Smriglio (1999) accepted to keep the two species separate, considering also that L. bedullii shows some very peculiar characteristics not recorded in other Leptochiton (e. g., the cylindrical dorsal spicules of the girdle), and we agree with this interpretation. Comparisons. The characteristics of the sculpture of Leptochiton boettgeri agree with the belonging of the species to the Leptochiton cancellatus group, see Tab. 2. The closest species is L. bedullii, as already reported by Dell’Angelo & Palazzi (1989).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA64E5A0FADFC8C6F1B931E.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Romania: Kostej (Šulc 1934); Czech Republic: Rudoltice (Šulc 1934); Poland: Korytnica, Węglinek (Bałuk 1971, 1984; Macioszczyk 1988).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	description	Fig. 30	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	materials_examined	Type material. Holotype MSNG 62633, intermediate valve, width 1.5 mm (Figs 30 E – G). Paratype 1, MSNG 62633, head valve from Rohožnik, width 1.5 mm (Figs 30 A – B). Paratype 2, SNM Z 38496 / a (as Leptochiton sp.), tail valve from Rohožnik, width 1.1 mm (Fig. 30 H). Type locality. Devínska Nová Ves (Slovenia). Type stage. Middle Miocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	etymology	Etymology. The name honors Andrej Ruman (Comenius University, Bratislava, Slovakia) for his contribution to the study of fossil mollusks.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	materials_examined	Material examined. Middle Miocene: Central Paratethys: Slovakia: Devínska Nová Ves brickyard: holotype plus 1 valve (BD 403; Rohožnik – konopiská clay pit (“ Hornáčková jama ”): paratype plus 6 valves (BD 404, Figs 30 C – D). Maximum width of valves: 1.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	diagnosis	Diagnosis. Valves fragile and small, head valve semicircular, intermediate valves broadly rectangular, rounded, elevated, apex inconspicuous, tail valve elliptical, mucro not prominent, in anterior position. Tegmentum surface rough, sculptured with roundish granules, arranged without pattern in HV, elliptical, narrow granules, concentrically arranged in LA, PMA, roundish granules, arranged without pattern, with generally one or two stems in apical part in CA, AMA, each granule with up to 4 – 5 aesthetes almost aligned in HV, LA, up to 5 – 7 aesthetes in CA, all more or less of same size.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	description	Description. Valves fragile and of small size. Head valve semicircular. Intermediate valves broadly rectangular (W / L = 1.91 – 2.18), rounded in anterior profile, elevated (H / W = 0.51 estimated), anterior margin straight, side margins rounded, posterior margin almost straight, apex inconspicuous, lateral area slightly raised, with few concentric growth marks. Tail valve elliptical, anterior margin convex, mucro not prominent, in anterior position, without any distinct diagonal ridge. Tegmentum surface rough, very dense but well separated granules. HV sculptured with roundish granules arranged without pattern, diameter 30 – 40 µm. LA and PMA sculptured with elliptical, narrow granules, concentrically arranged, long 30 – 40 µm. CA and AMA sculptured with roundish / polygonal granules very irregular, arranged without pattern, with generally one or two stems (or at least a hint of) in apical part, up to 65 µm long. Each granule with up to 4 – 5 aesthetes almost aligned in HV and LA, up to 5 – 7 aesthetes, both on top and peripheral margin in CA, all aesthetes more or less of same size. Articulamentum without insertion laminae, with small apophyses, widely separated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	discussion	Remarks. The fossil record of Leptochiton rumani sp. nov. is limited to the Middle Miocene of Slovakia. This species has already been reported and figured by Ruman & Hudáčková (2015), as Leptochiton sp. Despite the small size of the valves examined, often incomplete, the sculpture of the valves of Leptochiton rumani sp. nov. is different from that of the other species known from the Paratethys [(L. boettgeri (Šulc, 1934), L. sulci (Bałuk, 1971) and L. cancellatus (Sowerby, 1840)], all characterized by the presence of granules arranged in radial series on HV, LA and PMA, and in longitudinal series on CA and AMA (see above and tab. 4 b). Notwithstanding the incompletness of the holotype, we have estimated the dorsal elevation H / W (put in brackets in tab. 4). The granules have a very irregular structure, even aesthetes are hardly divisible in megalaesthetes and micraesthetes. In some granules the size of the pores of the aesthetes located centrally seem slightly larger than the others, but this could be due to erosion, for which we rate aesthetes more or less of the same size. Comparisons. The characteristics of the sculpture of Leptochiton rumani sp. nov. support its belonging to the Leptochiton tavianii group, see Tab. 4. However, all these species present a different tegmentum sculpture than the species here under study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA54E5C0FADF9F568A79490.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Slovakia: Devínska Nová Ves, Rohožník (Ruman & Hudáčková 2015; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA34E5D0FADFE6C694590D9.taxon	description	Fig. 31	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA34E5D0FADFE6C694590D9.taxon	description	Hudáčková 2015, p. 158, figs 2.2 – 2.6; Dell’Angelo et al. 2018 b, p. 52, tab. 17; Dulai 2025 a, p. 4, figs 3 – 8. Acanthochitona sp. Tomašových, 1998, p. 362, pl. 1, figs 7 – 8 (fide Ruman & Hudáčková 2015). non Leptochiton (Leptochiton) sulci; Studencka & Studencki 1988, p. 38, pl. 1, fig. 4 [= Leptochiton cancellatus (Sowerby, 1840), fide Studencka & Dulai 2010: 261).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA34E5D0FADFE6C694590D9.taxon	materials_examined	Type material. Holotype BkK-A 08 (Bałuk coll.), intermediate valve figured by Bałuk (1971: pl. 2, fig. 3). Type locality. Korytnica, 24 km SSW of Kielce, southern slopes of the Holy Cross Mts (Poland). Type stage. Middle Miocene. Material examined. No actual material available, only descriptions and illustrations from the literature (Figs 31 A – H). Maximum width of valves: 2.5 / 3.2 / 2.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA34E5D0FADFE6C694590D9.taxon	description	Description. Head valve semicircular. Intermediate valves broadly rectangular, rounded in anterior profile, moderately elevated, anterior and posterior margins almost straight, apex not indicated, lateral areas slightly raised, with few growth ridges. Tail valve semicircular, mucro in central position, postmucronal slope slightly concave. Tegmentum surface rough, granules disposed in longitudinal striae greater than those in radial ones. HV, LA and PMA sculptured with dense roundish granules arranged in radial striae (more than 50 in HV). CA and AMA sculptured with longitudinal series of roundish granules (CA 60 – 70, AMA ca. 50) joined by a narrow ridge, displaying a regular quincuncial pattern, with intercostal spaces twice lesser than their diameter. Each granule usually with 3 – 4 aesthetes of same size, normally aligned each other. Articulamentum without insertion laminae, apophyses narrow and widely spaced.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA34E5D0FADFE6C694590D9.taxon	discussion	Remarks. Leptochiton sulci (Bałuk, 1971) is very similar to L. cancellatus (Sowerby, 1840), a well known species from the Mediterranean area, and was considered conspecific with L. cancellatus first by Laghi (1977), and later also by other authors (Dell’Angelo & Palazzi 1989; Dell’Angelo & Smriglio 1999; Dell’Angelo & Silva 2003). Also WoRMS consider L. sulci as a synonym of L. cancellatus. However, the two taxa show differences discernable (Studencka & Dulai 2010; Ruman & Hudáčková 2015). In L. sulci, the granules are less dense, longitudinally arranged and joined by a narrow ridge, whereas in L. cancellatus they lay partially one upon the other; they differ also in the number and arrangement of the aesthetes: in L. cancellatus, each granule has a central megalaesthete with six lateral micraesthetes (Studencka & Dulai 2010: fig. 2 D), whereas there are only three-four aesthetes in L. sulci (Bałuk 1984: pl. 3, fig. 1 – 2; Ruman & Hudáčková 2015: fig. 2.4 b) (Figs 31 A – B, 31 F), normally aligned with each other and more or less of the same size, difficult to separate into micraesthetes and megalaesthetes aesthetes. Moreover, the granules of neighbouring longitudinal ribs of L. sulci display a regular “ quincuncial pattern ” (see Bałuk 1984: pl. 3, figs 1 – 2; Ruman & Hudáčková 2015: fig. 2.4 b) (Figs 31 A – B, 31 F), not present in L. cancellatus (see Kaas 1981: fig. 10 F; Studencka & Dulai 2010: fig. 2 D). Schematic sculptural differences for both these species are well illustrated by Macioszczyk (1988: fig. 3). So, we consider the two species as different, following Bałuk (1971, 1984), Studencka & Dulai (2010) and Ruman & Hudáčková (2015). Šulc (1934) described as Lepidopleurus cf. cancellatus (Capellini) a tail valve from Steinabrun in the Vienna Basin, not present at NHMW. The lack of figures does not permit to ascertain whether this valve truly pertains to Leptochiton cancellatus or instead to L. sulci, equally reported from the Parathetys. The material examined from Varovtsi and Horodok (Ukraine) includes many valves of a Leptochiton spp. which, after a thorough examination, do not agree with the attribution to L. boettgeri or L. sulci, but to L. cancellatus, corroborating the presence of this species from Varovtsi, as already reported by Studencka & Dulai (2010). Some of these valves are figured (see above, Figs 13 M – T). Comparisons. The characteristics of the sculpture of Leptochiton sulci agree with the belonging of the species to the Leptochiton cancellatus group, see Tab. 2.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA34E5D0FADFE6C694590D9.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Slovakia: Devínska Nová Ves, Rohožník (Tomašových 1998; Ruman & Hudáčková 2015; this study); Hungary: Devecser (Dulai 2025 a); Poland: Korytnica, Węglin (Bałuk 1971, 1984; Macioszczyk 1988).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA24E5F0FADF8B66E779490.taxon	description	Fig. 32	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA24E5F0FADF8B66E779490.taxon	materials_examined	Type locality. Höllkopf, near Glimmerode (Germany). Type stage. Oligocene (Chattian). Material examined. Oligocene (Chattian): Paratype SMF 250023 / 1 E, Figs 32 C – D (from R. Janssen, see Dell’Angelo & Palazzi 1989: 76). Maximum width of the valves: 4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA24E5F0FADF8B66E779490.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, slope straight. Intermediate valves broadly rectangular, width more than three times the length, flat, rounded in anterior profile, anterior margin almost straight, side margins rounded, posterior margin straight, apex not developed, lateral areas not raised. Tail valve semicircular (W / L = 1.86 – 1.95), flat, anterior margin straight, mucro slightly protruding, in anterior position. Tegmentum with growth lines clearly marked. HV, LA and PMA sculptured with radial series of dense, minute, roundish granules (LA 10 – 13, PMA 63 – 68). CA and AMA sculptured with 16 – 20 longitudinal series of granules barely visible. Articulamentum lacking insertion laminae to girdle, with large and triangular apophyses.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA24E5F0FADF8B66E779490.taxon	discussion	Remarks. Leptochiton chatticus (Janssen, 1978) is only known for the original description, based on the three types of valves (head, intermediate and tail), 15 in total. It shows, according to the author, remarkable affinities with L. maguntiacus de Rochebrune, 1882, from which it differs to be flatter, for the more elliptical shape of the tail valve and the mucro less protruding and more displaced anteriorly. Unfortunately, the original description is rather short, so as to leave many uncertainties, and the iconography presented is insufficient. Fortunately, Janssen sent a paratype to the senior author, illustrated in the drawing, Figs 32 C – D (Dell’Angelo & Palazzi 1989: 77), which made possible to better define some characteristics of the species. Gürs (1995) did not report Leptochiton chatticus in his Thesis, considering the paratype SMF 250022 as a juvenile specimen of Lepidopleurus virgifer (Sandberger, 1859) (Gürs 1995, p. 19: “ in juvenile individuals the granules not yet fused into radial ribs ”. Gürs (1995) described in his Thesis a new species (Lepidopleurus jansseni) from the lower Oligocene of Eckelsheim (Germany), based on the three valves. The thesis was not published, and the taxon Lepidopleurus jansseni was not formalized. The material of Lepidopleurus jansseni, not present at SMF (R. Janssen, pers. comm., 2020) is possibly kept in the Gürs collection. The “ type ” material (with the three types of valves) was figured by Gürs (1995) and we report here these same illustrations (Figs 32 E – H). Based on the descriptions and iconography, Lepidopleurus jansseni is morphologically indistinguishable from L. chatticus, and we prefer therefore to synonymize the two taxa. Comparisons. The characteristics of the sculpture of Leptochiton chatticus Janssen, 1978 agree with the belonging of the species to the Leptochiton cancellatus group, see Tab. 2. The species is strongly depressed, which differentiates it from L. cancellatus (Sowerby, 1840) and L. sarsi Kaas, 1981, more similar for the shape and size of the granules. Very characteristic is the tail valve, whose mucro is very displaced anteriorly, so that the antemucronal area is reduced to a narrow strip with longitudinal striae of granules formed by a few coarse granules (from 2 to 6) which tend to coalesce.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA24E5F0FADF8B66E779490.taxon	distribution	Distribution. Lower Oligocene (Rupelian): North Europe: Germany: Steigerberg (Gürs 1995). Upper Oligocene (Chattian): North Europe: Germany: Glimmerode (Janssen 1978).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA04E400FADFE6C6E769628.taxon	description	Fig. 33	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA04E400FADFE6C6E769628.taxon	materials_examined	Type material. Holotype, tail valve, figured by Gürs (1992: pl. 1, fig. 1 d), Fig. 33 B. Paratypes: SMF, more than 250 valves, figured by Gürs (1992: pl. 1, fig. 1 a – 1 c, 1 e), Figs 33 A, 33 C, 33 E – H. Holotype of Lepidopleurus paeninsulae, tail valve, figured by Gürs (1992: pl. 1, fig. 2 b), Fig. 33 D. Type locality. Steigerberg, near Eckelsheim (Germany). Type stage. Oligocene (Rupelian). Material examined. No actual material available, only descriptions and illustrations from the literature.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA04E400FADFE6C6E769628.taxon	description	Description. Head valve semicircular, flat, posterior margin widely V-shaped. Intermediate valve broadly rectangular, narrow, moderately elevated (H / W = 0.35), semicarinate in anterior profile, posterior margin straight, apices not developed, lateral areas raised. Tail valve semicircular, mucro in anterior position, antemucronal area very narrow. Tegmentum rough. HV, LA and PMA sculptured with radial granular striae intersected by concentric growth lines, with some elevated pustules irregularly distributed on striae, concentric growth lines continuing across central area, where they weaken, becoming inconspicuous. CA and AMA sculptured with short longitudinal striae of roundish granules, weakening anteriorly, subobsolete in jugal area, which appears smooth. Articulamentum without insertion laminae, apophyses well developed, bluntly triangular with a rounded margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA04E400FADFE6C6E769628.taxon	discussion	Remarks. The description given by Gürs (1992) is incomplete, as important diagnostic information is missing. Moreover, the two intermediate and the two tail valves figured seem to differ in shape (compare Gürs 1992, figs 1 b – c and 1 d – e). The intermediate valve in fig. 1 b could be interpreted as a valve ii (often a bit different from other intermediate valves iii – vii) and both of the valves have the longitudinal granular striae on pleural areas short, whereas Gürs (1992) noted in his description that in some other intermediate valves the sculpture may reach the anterior valve margin. Gürs (1992) described a second species (Lepidopleurus peninsulae) from the Oligocene (Chattian) of Höllkopf (Germany), that differs by the coarser pustular sculpture on the lateral areas, the smaller size of the valves and fewer radial granular striae on the postmucronal area. We do not consider these differences sufficient to warrant the specific validity of this species, and we consider L. paeninsulae a synonym of L. eckelsheimensis: this is agreement with Gürs (1995) who report Lepidopleurus peninsulae only in the discussion of Lepidopleurus jansseni. Comparisons. The characteristics of the sculpture of Leptochiton eckelsheimensis (Gürs, 1992) agree with the Leptochiton cancellatus group (see Tab. 2). Leptochiton eckelsheimensis is rather similar to L. lateropustulosus Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 from the Miocene (Tortonian) of Saint-Clément-de-la-Place (France) (see below), from which it differs mainly for the conspicuous difference in stratigraphic age, and the unavailability of the type material of L. eckelsheimensis for more accurate comparisons.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFA04E400FADFE6C6E769628.taxon	distribution	Distribution. Lower Oligocene (Rupelian): North Europe: Germany: Steigerberg (Gürs 1995). Upper Oligocene (Chattian): North Europe: Germany: Höllkopf (Gürs 1992, 1995).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBF4E410FADFCF469CA9235.taxon	description	Fig. 34	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBF4E410FADFCF469CA9235.taxon	materials_examined	Type material. SMF, Lectotype (intermediate valve) and paralectotype (head valve) designated by Janssen (1978: 219). Type locality. Gienberg (Germany). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Germany: Steigerberg: 10 valves (BD 405, Figs 34 A – L). Maximum width of the valves: 4.8 / 8 / 7.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBF4E410FADFCF469CA9235.taxon	description	Description. Head valve less than semicircular, posterior margin widely V-shaped, notched in middle. Intermediate valves broadly rectangular, elongate (W / L = 2.8), semicarinate in anterior profile, moderately elevated (H / W = 0.35 – 0.41), anterior margin straight, side margins rounded, posterior margin straight, apex not indicated, lateral areas hardly raised, though clearly indicated. Tail valve semicircular (W / L = 1.86), front margin straight, mucro subcentral, antemucronal slope slightly convex, postmucronal slope concave. Tegmentum rough, growth lines are often present in variable numbers. HV, LA, PMA sculptured with dense, minute, roundish granules arranged in radial series (70 in HV, 60 – 70 in PMA), initially fine, becoming coarser towards the margins. CA, AMA sculptured with roundish granules (diameter 43 – 57 µm) arranged in longitudinal series (CA 60 – 65, AMA 60 – 70 and joined together, space between striae of granules large. Each granule with a central megalaesthete and up to 8 micraesthetes around edge. Articulamentum lacking insertion laminae to the girdle, apophyses small, widely spaced, triangular but tending to become rounded in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBF4E410FADFCF469CA9235.taxon	discussion	Remarks. Leptochiton maguntiacus (de Rochebrune 1882) was always classified in the old collections as Lepidopleurus virgifer. A thorough discussion given by past authors (including Boettger) is reported in Janssen (1978: 220). Boettger was the first to recognize that in addition to Lepidopleurus virgifer even finer sculpted forms occur, and called this material Chiton virgifer? juv. var. agnata, but without publishing this taxon. Dell’Angelo & Palazzi (1989: 64) considered Leptochiton maguntiacus as a synonym of L. algesirensis (Capellini, 1859). The two species undoubtedly have remarkable similarities, well highlighted by Dell’Angelo & Palazzi (1989), but the rougher sculpture of L. maguntiacus, especially in HV, LA and PMA (minute, roundish granules arranged in radial series, initially fine, becoming coarser towards the margins) is different from that of L. algesirensis (more regular radial series), and this alone is sufficient to separate the two species. However, the two species differ in other features (e. g., the larger space between striae of granules in CA and AMA, the anterior profile of the intermediate valves more rounded and a lower dorsal elevation in L. algesirensis) and in the stratigraphic distribution [L. algesirensis does not exceed the Miocene (Tortonian)]. Comparisons. The characteristics of the sculpture of Leptochiton maguntiacus agree with the belonging of the species to the Leptochiton cancellatus group.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBF4E410FADFCF469CA9235.taxon	distribution	Distribution. Lower Oligocene: North Europe, Germany: Eckelsheim, Gienberg, Heimberg, Trift, Zeilstück, Welschberg, Würzmühle (Janssen 1978; Gürs 1983, 1995; Hocht 1986).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBE4E430FADF8C868869794.taxon	description	Fig. 35	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBE4E430FADF8C868869794.taxon	description	Lepidopleurus aff. poirieri; Müller 2011, p. 21, pl. 4, figs 1 – 6.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBE4E430FADF8C868869794.taxon	materials_examined	Type material. Lectotype, designated by Janssen 1978, tail valve figured by de Rochebrune (1882: pl. 1, fig. 10 b). Type locality. Jeurre, near Étampes (France). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene: France: Gaas (Espibos, Lagouarde): 11 valves (AC, DA, Figs 35 H – I, PR). Upper Oligocene: Germany: Glimmerode: 8 valves (BD 256, Figs 35 A – G, 35 J – L). Maximum width of the valves: 3.6 / 2.7 / 3.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBE4E430FADF8C868869794.taxon	description	Description: Head valve semicircular, slope straight. Intermediate valves rectangular (W / L = 2.50 – 2.53), rounded in anterior profile, moderately elevated (H / W = 0.32 – 0.43), anterior margin straight, lateral margins slightly rounded, posterior margin straight, apex not indicated, lateral areas little raised. Tail valve semicircular (W / L = 1.86 – 2.00), anterior margin straight, mucro in central position, antemucronal slope straight or slightly convex, postmucronal slope almost slightly concave. Tegmentum smooth, growth lines often present in variable numbers. HV, LA and PMA sculptured with radial series of dense, minute, roundish granules (HV 90 – 100, LA 20 – 22, PMA 75 – 80) joined together. CA and AMA sculptured with longitudinal series of more elliptical granules, partially overlapping (CA 80 – 85), with smooth and large intercostal spaces. Each granule usually with 1 megalaesthete in central position and up to 6 micraesthetes along margin. Articulamentum lacking insertion laminae to the girdle. Apophyses are small, widely spaced, triangular but tending to become rounded in tail valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBE4E430FADF8C868869794.taxon	discussion	Remarks. This species was described on material from localities of French Oligocene, Jeurre and Étréchy (de Rochebrune 1882; Cossmann 1892) and was confirmed by Janssen (1978) for the French middle Oligocene (Auvers-Saint-Georges) and for the German middle to late Oligocene. Comparisons. The characteristics of the sculpture of Leptochiton poirieri (de Rochebrune 1882) agree with the belonging of the species to the Leptochiton cancellatus group, see Tab. 2. Many Leptochiton species have a similar tegmental sculpture, two of them [L. maguntiacus (de Rochebrune, 1882) and L. chatticus (Janssen, 1978)] are known from the Oligocene of Germany, the others still extant in Mediterranean Sea or Atlantic Ocean, and / or with a stratigraphic distribution that does not extend deeper than the Miocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBE4E430FADF8C868869794.taxon	distribution	Distribution. Lower Oligocene: North Europe, Germany: Eckelsheim, Gienberg, Trift, Welschberg, Zeilstück, (Janssen 1978; Hocht 1986; Gürs 1995; Welle 2009; Müller 2011); Northeastern Atlantic (Rupelian): France: Paris Basin: Auvers-Saint-Georges (Janssen 1978), Etrechy (de Rochebrune 1882); Aquitaine Basin: Gaas, Jeurre (Dell’Angelo et al. 2018 a; Cherns & Schwabe 2019); Upper Oligocene: North Europe, Germany: Glimmerode, Freden, Söllingen, Doberg, Rummeln (Janssen 1978).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	description	Fig. 36	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	materials_examined	Type material. Lectotype SMF 250010, designated by Janssen (1978), tail valve figured by Koenen (1892: figs 24 a – c) and Janssen (1978: pl. 14, fig. 1), Fig. 36 A; Paralectotype, tail valve figured by Koenen (1892: figs 25 a – b). Type locality. Brandhorst (Westphalia, Germany). Type stage. Lower Oligocene (Rupelian). Material examined. No actual material available, only descriptions and illustrations from the literature.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valve broadly rectangular, anterior margin straight, rounded in anterior profile, side margins slightly rounded, posterior margin straight, apex not developed, lateral areas not raised, barely differentiated from central area. Tail valve semicircular, anterior margin straight, mucro in central position, just prominent. HV and PMA sculptured with radial striae of roundish granules on head valve and postmucronal area of tail valve, LA finely granulated such as to appear almost smooth. CA and AMA sculptured with longitudinal granular striae of roundish granules. Articulamentum without insertion laminae, apophyses small, widely spaced.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	discussion	Remarks. The descriptions by Koenen (1892) and Janssen (1978) are based only on intermediate and tail valves. Gürs (1995: pl. 1, fig. 5 – 7) illustrates three valves (Figs 36 B – D), also including a head valve (Fig. 36 B). Comparisons. The characteristics of the sculpture of Leptochiton uhligi (Koenen, 1892) agree with the belonging of the species to the Leptochiton cancellatus group, see Tab. 2, albeit with some doubts. The sculpture of lateral areas of intermediate valves is not so clear seeing Fig. 36 C, Janssen (1978) reports areas finely granulated such as to appear almost smooth, and this raises some doubts about the presence of radial striae of granules. Leptochiton uhligi differs from other Leptochiton species by the particularly pronounced central area, the lateral areas barely differentiated, and the mucro of the tail valve just prominent (Janssen 1978).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	distribution	Distribution. Lower Oligocene: North Europe, Germany: Böseberg, Brandhorst (Janssen 1978). Genus Belknapchiton Sirenko, Saito & Schwabe, 2022	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	type_taxon	Type species. Leptochiton belknapi Dall, 1878 by original designation.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	distribution	Distribution. Belknapchiton is known from the Pliocene to the Recent. To date there are 22 Recent species known, the vast majority of which inhabit the deep waters of the Pacific Ocean and only B. alveolus (M. Sars MS, Lovén, 1846) is known from the Atlantic Ocean. The fossil record includes the Pliocene to Pleistocene of Italy (Dell’Angelo et al. 2013, 2021 b; Taviani et al. 2023).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBC4E440FADFD686978912C.taxon	discussion	Remarks. The genus Belknapchiton was recently established by Sirenko et al. (2022), and contains a widespread group of species conventionally assigned to Leptochiton. The new genus differs from other representatives of the family Leptochitonidae by having an elongate oval shape of body, small rather narrow, sharply pointed scales and scattered long, smooth, needles, the combination of a wide central tooth with short first lateral teeth, and large head of major lateral teeth of radula occasionally with an additional small denticle-like appendage at inner edge. The main morphological characters of Belknapchiton alveolus are reported in Tab. 5, for ease of comparison with other spp. of Leptochiton showing similar features.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	description	Fig. 37	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	materials_examined	Type material. Lectotype NRS, type collection n. 104, one specimen in alcohol, the valves missing (designed by Kaas 1981: 223). Type locality. Bohuslän (Sweden). Material examined. Lower Pliocene: Italy: Borzoli: 6 valves (BD 407, MSNG); Zinola: 1 valve (MZB 45701, Figs 37 A – B). Pleistocene, presumably last glacial: Italy: off Bari, cruise SE 06: SE 06 - 35: 1 valve; SE 06 - 40: 18 valves (Figs 37 E – G); SE 06 - 50: 56 valves (Fig. 37 H). Recent. Norway, Sula Ridge, dredged at 215 m (MZB, Figs 37 C – D). Maximum width of the valves: 4.3 / 5.7 / 3.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	description	Description. Valves elevate and arcuate. Head valve semicircular, posterior margin widely V-shaped, notched in middle, slope straight. Intermediate valves broadly rectangular (W / L = 1.78 – 2.01), arched in anterior profile, elevate (H / W = 0.46), anterior margin straight, side margins rounded, posterior margin almost straight, without a distinct apex, lateral area indiscernible from central area, with few concentric growth marks. Tail valve more than semicircular (W / L = 1.38 – 1.66), anterior margin almost straight in wide jugal area, mucro not prominent, posteriorly placed at ⅓ from posterior margin, postmucronal slope concave. Tegmentum rough, surface densely covered with oval microgranules disposing in quincunx, granules (length 70 – 80 µm) with one posterior megalaesthete and 4 – 5 micresthetes upward. Articulamentum without insertion laminae, with small apophyses, sharply triangular, widely separated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	discussion	Remarks. Belknapchiton alveolus (M. Sars MS, Lovén, 1846) was originally described from the west coast of Scandinavia (Bohuslän, Sweden, and Bergen, Norway), but the species was reported under a dozen different names from elsewhere, and was initially considered as the only cosmopolitan species occurring in three oceans (Kaas & Van Belle 1985 a). Recently Wu & Okutani (1984) have demonstrated that B. belknapi (Dall, 1878), which is closely related to B. alveolus, shows several differential characters which warrant a specific separation, and a study of the geographical distribution of the synonymous species indicates that B. alveolus seems to be restricted to the Atlantic Ocean, while B. belknapi is confined to the Pacific and lndian Oceans. A good illustration of the valves of Recent specimens of Leptochiton alveolus is given by Wu & Okutani (1984). Many valves of Belknapchiton alveolus from Sea-bottom sampling at bathyal depths off the Apulian margin (southwestern Adriatic Sea, SE 06 cruise) have recently been studied by Taviani et al. (2023). The studied material (Figs 37 E – H) fully agree with the attribution to B. alveolus, compare with the figures of recent valves from Japan (Wu & Okutani 1984: pl. 3, figs 9 – 14; pl. 4, fig. 1) and a single intermediate valve from Sula Ridge, Norway, dredged at 215 m (Figs 37 C – D). The finding of B. alveolus from the SE 06 cruise is quite significant. This taxon is now extinct in the Mediterranean basin, also if some older reports from this area were erroneously recorded, i. e. Vayssiere (1913: 32 « Napoli, Palermo, Dalmatia », probably on the basis of records from previous authors) and Van Belle (1975), two specimens collected by fishermen at 200 - 250 m from Gerona, Spain, but the identification was proven wrong, and they have been reclassified as Leptochiton geronensis Kaas & Van Belle, 1985, by the same authors. Specimens reported as “ Leptochiton alveolus ” from Eocene-Oligocene cold-seep limestones of the Olympic Peninsula, Washington (Squires & Goedert 1995; Peckmann et al. 2002, Schwabe & Sellanes 2010) has not been considered in the distribution of B. alveolus, because their specific and even generic assignment remains an open problem (Schwabe & Sellanes 2010; Hybertsen & Kiel 2018). Comparisons. See Tab. 5 for a comparison with the species of the Leptochiton tavianii group considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	distribution	Distribution. Lower Pliocene: Italy, Liguria: Borzoli, Zinola (Dell’Angelo et al. 2013, 2021 b). Pleistocene, presumably last glacial: Italy: off Bari (Taviani et al. 2023). Recent: North Atlantic: all along the Norwegian and Swedish west coast (Kaas 1981; Hansson 1988), Iceland (Sneli & Gudmundsson 2018), Bay of Biscay (Kaas 1979), Spain, Galicia (Rolan Mosquera et al. 1990; Urgorri et al. 2017), NW Portugal (Consolado Macedo et al. 1999), Gulf of Maine and Gulf of St. Lawrence (Wu & Okutani 1984). Genus Parachiton Thiele, 1909	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	type_taxon	Type species. Lepidopleurus (Parachiton) acuminatus Thiele, 1909 by original designation.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	distribution	Distribution. Parachiton is known from the Miocene to the Recent. To date there are 23 Recent species known, all from the Indo-West Pacific except for P. africanus (Nierstrasz, 1906) from the Mediterranean Sea. The fossil record includes the lower Miocene of France (Dell’Angelo et al. 2018 a), the Middle Miocene of Paratethys (Šulc 1934; Bałuk 1984; Ruman & Hudácková 2015), the Miocene to Pleistocene of Italy (Dell’Angelo et al. 2013, 2015 a, 2018 a) and the Pleistocene deposits of the Red Sea (Dell’Angelo et al. 2020 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFBB4E460FADFBC96B5A96F5.taxon	discussion	Remarks. Parachiton was originally established by Thiele (1909) as a subgenus of Lepidopleurus Risso, 1826 on the basis of its disproportionately large tail valve with subterminal mucro, and overall similarities of the other valves with species of Lepidopleurus. It is now considered to be a distinct genus (Saito 1996; Sirenko 2006), also due to differences in the radula (Saito 1996). The sculpture is characteristic, with rows of granules with 3 pores of aesthetes, arranged usually longitudinally in the central area of the intermediate valves and also in the antemucronal area of tail valve. The main morphological characters of the Parachiton spp. considered in the present study are reported in Tab. 6. observations).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB84E490FADFDCA6F3893A4.taxon	description	Fig. 38	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB84E490FADFDCA6F3893A4.taxon	materials_examined	Type material. Holotype: RMNH, Leiden, no. reg. 2783, a specimen deprived of its end valves (Kaas 1977). Holotype of Parachiton thielei figured by Šulc 1934 (Fig. 38 I). Type locality. Oran, Algeria. Material examined. Middle Miocene: Central Paratethys: Austria: Steinabrunn: 1 valve (NHMV 1933 / 0001 / 0042, Figs 38 M – P; Hungary: Letkés: 1 valve (BD 408). Pliocene: Spain: Estepona: 1 valve (BD 409). Italy: Piedmont: Valle Andona: 1 valve (BD 410); Emilia-Romagna: Cava di Campore: 3 valves (BD 411). Pleistocene: Italy: Calabria: Archi S. Francesco: 9 valves (BD 412, Figs 38 A – F), Carrabbati: 1 valve (BD 413), Gallina: 1 valve (BD 414, Figs 38 G – H), Petti di Carrubbare: 2 valves (BD 415), Terreti: 3 valves (BD 416); Sicily: Capo Milazzo: 1 valve (BD 417). Maximum width of the valves: 5 / 7 / 5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB84E490FADFDCA6F3893A4.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.09 – 2.18), rounded in anterior profile, moderately elevated (H / W = 0.30 – 0.36), anterior margin straight between the apophyses, posterior margin straight, apex not indicated, lateral areas only discernible by difference in sculpture, marked with 6 or 7 faint concentric lines of growth. Tail valve from elliptical to almost triangular (W / L = 1.14 – 1.28), mucro somewhat swollen, rounded, posterior, at 7 / 8 of valve’s length, antemucronal slope slightly convex, postmucronal slope short, steep, straight, but for a little excavation directly behind the mucro. Tegmentum finely granulose all over. HV and PMA sculptured with radial series of more or less oval granules, interrupted by weak growth lines. LA sculptured with granules quincuncially arranged, tending towards an arrangement in radial striae, interrupted by 6 – 7 concentric growth lines. CA and AMA sculptured with longitudinal series of granules (CA 45 – 55, AMA 38 – 50), with reduced space between striae, arranged side by side, with exception of jugal tract, where they are slightly convergent, on sides sometimes anastomosing, granules more clearly pronounced towards side margins. Articulamentum weakly developed, apophyses small, more or less triangular, wide apart, jugal sinus broad, slightly convex or flat.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB84E490FADFDCA6F3893A4.taxon	discussion	Remarks. Parachiton africanus (Nierstrasz, 1906) is a rare Mediterranean species described upon a single specimen of small dimensions (length 7 mm, width 3 mm), from Oran, Algeria. Based upon a second specimen found after 70 years later at Gallipoli (Puglia, Italy) compared with Nierstrasz’s holotype, Kaas (1977) validates the taxon, thus far apparently endemic to the Mediterranean. The fossil species Lepidopleurus (Parachiton) thielei was described by Šulc (1934) on the basis of a few intermediate and tail valves from the Miocene deposits of Steinabrunn in the Vienna Basin, and afterwards from Korytnica (Poland) by Bałuk (1971, 1984). Kaas (1977) stated: “ in my opinion L. (P.) africanus, if not conspecific with L. (P.) thielei, must be regarded as a closely allied descendant of the latter ”). It was considered as a synonym of Parachiton africanus by several authors (Laghi et al. 1981; Bałuk 1984; Kaas & Van Belle 1985 a; Dell’Angelo & Palazzi 1989; Dell’Angelo et al. 2015 a, 2018 a). The only differences pointed out are those reported by Bałuk (1984: “ the differences concern only greater dimensions and an absence of regular interspaces between the concentric ridges on lateral areas of intermediate valves in L. thielei ”). In order to explore whether there is ground to discriminate between the two taxa under scrutiny upon morphological arguments, we have analyzed in-depth the following features: The tegmentum’s sculpture in CA and AMA. Nierstrasz (1906) does not report the number of longitudinal rows for Parachiton africanus, he says “ at least 36 ” for intermediate valves, and “ sculptured like the central area of the middle valves ” for tail valve. It is difficult to count the number of longitudinal striae of granules in CA and AMA, the striae are mainly parallel and regular but sometimes anastomosing and often converging posteriorly on the jugum, and by no means as regular as Nierstrasz’s figures would suggest. Šulc reports ca. 50 longitudinal rows of granules for Parachiton thielei, parallel except on the jugal tract, where they converge (not in Šulc’s description, but evident in his fig. 4 of the holotype). In the valve at NHMW the longitudinal ribbing on the jugal tract is a little convergent indeed, and the number of longitudinal rows of granules agree with the 50 reported by Šulc. A study of all intermediate and tail fossil valves available and some recent ones allow us to better define the range of radial striae of granules in CA (45 – 55) and AMA (38 – 50), so also for this feature, a certain degree of variability seem acceptable. The shape of tail valves. The shape of tail valves of Parachiton africanus is variable, from an elliptical, almost triangular profile, as in the holotype figured in Nierstraz (1906: figs 1, 6) to a more regularly rounded profile, as in the recent valves figured by Kaas (1977: figs 1, 6) and Dell’Angelo & Smriglio (1999: pl. 23, fig. F). The same variability is present in the fossil valves object of this study (compare Fig. 38 E and Fig. 38 G). The same variability can be observed in Parachiton thielei. Šulc figured two tail valves (Figs 38 I – J), of which the first, larger (W = 3.8 mm), designated as holotype, shows an elliptical, almost triangular profile, while the second one, smaller, is more regularly rounded, which could suggest a range of variability of the tail valve’s shape, as already suggested by Šulc (1934: “ On the basis of these tail valves it can be deduced that in immature individuals they do not have such an elliptical shape, but a more circular one ”). However, the tail valve present in the Šulc collection at NHMW (Figs 38 M – P) has the margin of PMA more regularly rounded and has a wide (3 mm) more like that of the holotype (3.8 mm), not in full agreement with the previously hypothesized range of variability of the tail valve profile. The possibility P. thielei being the ancestor of the recent P. africanus is still possible, or even likely. However, we did not find robust morphological evidence to separate the two species and this option would rest solely on biogeographic and paleoclimatic arguments. P. thielei inhabited subtropical contexts of the European Neogene vs. the temperate habitats of the Pleistocene to Recent P. africanus. Therefore, in consideration of the high degree of variability found on the examined material, we prefer to consider Parachiton thielei as a synonym of P. africanus. Comparisons. See Tab. 6 for a comparison with the Parachiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB84E490FADFDCA6F3893A4.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria: Steinabrunn (Šulc 1934; this study); Slovakia: Rohožník (Ruman & Hudácková 2015); Poland: Korytnica (Bałuk 1971, 1984); Hungary: Devecser, Letkés ¸ Várpalota (Dulai & Katona 2024; Dulai 2025 a, 2025 b; this study). Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Valle Andona (Laghi et al. 1981), Castell’Arquato (Dell’Angelo & Palazzi 1989), Cava di Campore (this study), Colle Sabbaco (Mancini 1998, 1999). Pleistocene: central Mediterranean, S. Italy: Arangea (Dell’Angelo & Palazzi 1989), Archi (Crovato & Taviani 1985), Terreti (Dell’Angelo & Smriglio 1999), Carrabbati, Gallina, Petti di Carrubbare, Capo Milazzo (this study). Recent: Northern part of the Atlantic coast of Morocco (Kaas et al. 2006), and the Mediterranean Sea: Spain (Costa Brava), France (Corsica), Italy: Gulf of Taranto, Strait of Sicily, Pantelleria island (Dell’Angelo et al. 1998 a), Turkey (Ozturk et al. 2014), Lebanon (Crocetta et al. 2014), Algeria: Oran (Dell’Angelo & Smriglio 1999).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB64E4A0FADF9786BBC90FB.taxon	description	Fig. 39	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB64E4A0FADF9786BBC90FB.taxon	materials_examined	Type material. Holotype: MHNBx 2017.9.1, tail valve, width 3.2 mm (Figs 39 A – C). Paratype: MZB 32134, tail valve, width 3 mm (Fig. 39 D). Type locality. Carrière Vives (France). Type stage. Miocene (Burdigalian). Material examined. Lower Miocene: France (Burdigalian): type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB64E4A0FADF9786BBC90FB.taxon	description	Description. Head and intermediate valves unknown. Tail valve semicircular, wider than long (W / L estimated 1.66), depressed, rounded in anterior profile, mucro posterior, located at about 2 / 10 th of valve’s length towards posterior margin, antemucronal slope slightly convex, postmucronal slope straight, but for a little excavation directly behind mucro. Tegmentum finely granulose. AMA sculptured with oval fine granules arranged in 50 enough regular longitudinal series, diverging anteriorly on sides. PMA sculptured with roundish granules arranged in ca 70 radial series, crossed by two concentric growth lines. Articulamentum weakly developed, apophyses small.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB64E4A0FADF9786BBC90FB.taxon	discussion	Remarks. Parachiton palmorum Dell’Angelo, Lesport, Cluzaud & Sosso, 2018 is only known from the original description, based on two tail valves from the lower Miocene of Carrière Vives (France). Comparisons. This species resembles Parachiton africanus (Nierstrasz, 1906), from which differs by the shape of the tail valve (much wider than long in P. palmorum, W / L estimated ca. 1.66 vs. 1.14 – 1.28 in P. africanus), the position of the mucro (located more posteriorly in P. africanus), and the sculpture of the longitudinal rows of granules, well evidenced and not coalescing in P. palmorum, fairly smoothly coalescing in P. africanus. Parachiton palmorum differs from P. statianus Dell’Angelo, Giuntelli, Sosso & Zunino, 2015 by the position of the mucro (located more posteriorly in P. statianus), the sculpture of the longitudinal rows of granules (well evidenced and not coalescing in P. palmorum, fairly smoothly coalescing in P. statianus), and the number of longitudinal (50 vs. 90 – 95) and radial (70 vs.> 100) rows of granules.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB64E4A0FADF9786BBC90FB.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Carrière Vives (Dell’Angelo et al. 2018 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB54E4B0FADF985695293DA.taxon	description	(Fig. 40) Lepidopleurus (Parachiton) aff. africanus [non Parachiton africanus (Nierstraz, 1906)]; Dell’Angelo et al. 2004, p. 29, pl. 2, figs 2, 6 (partim, fide Dell’Angelo et al. 2015 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB54E4B0FADF985695293DA.taxon	materials_examined	Type material. Holotype: MGPT PU 108784, tail valve, width 5.7 mm (Figs 40 G – H). Paratypes: MGPT PU 108785, intermediate valve (Figs 40 E – F); NHMW 2014 / 0450 / 0001 – 0002, intermediate and tail valve); BD 122 – BD 123, intermediate and tail valves. Type locality. Rio di Bocca d’Asino, Alessandria (Piedmont, Italy). Type stage. Upper Miocene (Tortonian). Material examined. Upper Miocene: Italy (Tortonian): type material plus Rio di Bocca d’Asino: 1 valve (MZB 32047). France (Messinian?), Ligerian Basin: Moulin-Pochas: 1 valve (PR, Fig. 40 D). Pliocene: Spain: Estepona: 2 valves (BD 418, MZB 32048, Fig. 40 C). Pleistocene: Italy: Sicily: Capo Milazzo: 1 valve (BD 419, Figs 40 A – B). Maximum width of the valves: -- / 6 / 5.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB54E4B0FADF985695293DA.taxon	description	Description. Head valve unknown. Intermediate valves broadly rectangular, rounded in anterior profile, moderately elevated (H / W = 0.35 – 0.40), anterior and posterior margins straight, lateral areas not raised. Tail valve semicircular (W / L = 1.51), anterior margin almost straight, mucro posterior, antemucronal slope slightly convex, postmucronal slope short, steep, straight. Tegmentum finely granulose. LA sculptured with not very evident irregularly arranged low granules, crossed by concentric lines of growth, that give a rough pattern to surface. PMA sculptured with more squarish granules arranged in numerous fine radiating rows, more than 100 in holotype. CA and AMA sculptured with longitudinal series of granules smoothly coalescing (CA 90 or more; AMA 95), rows enough regular, diverging anteriorly on sides. Articulamentum weakly developed, small apophyses.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB54E4B0FADF985695293DA.taxon	discussion	Remarks. Parachiton statianus Dell’Angelo, Giuntelli, Sosso & Zunino, 2015 is a rare species recently described on few valves for most incomplete, only the holotype is represented by a complete tail valve. The fossil record is known from the upper Miocene of the Ligerian Basin (France) and Po Basin (N. Italy), the Pliocene of Estepona Basin (Spain) and the Pleistocene of S. Italy. Comparisons. This species closely resembles Parachiton africanus (Nierstrasz, 1906).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB54E4B0FADF985695293DA.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic: Ligerian Basin, France: Moulin Pochas (Dell’Angelo et al. 2018 a); Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Rio di Bocca d’Asino (Dell’Angelo et al. 2015 a). Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004). Pleistocene: central Mediterranean, S. Italy: Capo Milazzo (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB44E4F0FADF8AA69FE9724.taxon	type_taxon	Type species. Hanleya debilis Gray, 1857 (= Chiton hanleyi Bean in Thorpe, 1844), by monotypy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB44E4F0FADF8AA69FE9724.taxon	distribution	Distribution. The genus Hanleya is recorded from the lower Oligocene onwards. Currently, all known recent species of Hanleya occur only in the Atlantic Ocean and adjacent seas, from off Brazil (25.44 ° S) to the Barents Sea (74.27 ° N) (Sirenko et al. 2016). The fossil record includes the Oligocene of Germany (Janssen 1978), the Middle Miocene of Paratethys (Šulc 1934; Ruman & Hudácková 2015), the upper Miocene of France (Dell’Angelo et al. 2018 b), the upper Miocene to the Pleistocene of Europe (Marquet 1984, 2002; Dell’Angelo et al. 2015 a, 2018 b, 2024; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB44E4F0FADF8AA69FE9724.taxon	discussion	Remarks. Revision of recent Hanleya recorded to date in the Atlantic Ocean and Mediterranean Sea is provided by Sirenko (2014) and Sirenko et al. (2016), with two species described as new, one from the Atlantic (H. harasewychi Sirenko, 2014) and one from the Mediterranean (H. mediterranea Sirenko, 2014). These authors considered H. nagelfar (Lovén, 1846) to be a junior synonym of H. hanleyi. The morphological characters of Hanleya spp. considered in the present study are reported in Tab. 7.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	description	Fig. 41	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	materials_examined	Type material. Holotype MSNG 62636, intermediate valve, width 4.6 mm (Figs 41 E – H, from Salice (Italy). Paratype 1, MSNG 62637, head valve, width 6.1 mm (Fig. 41 A), from Pezzo (Italy). Paratype 2, MSNG 62638, tail valve, width 5.4 mm (Figs 41 I – L), from Vallone Catrica (Italy). Type locality. Salice (Sicily, Italy). Type stage. Lower Pleistocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	etymology	Etymology. The name honors Giorgio and Pierluigi Taviani, elder brothers of M. T., for their continuous support and encouragement to pursue his vocation for marine sciences and paleontology; from the genitive of fratres (Latin for brothers).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	materials_examined	Material examined. Pleistocene: Italy: type material plus Calabria: Musalà: 2 valves (BD 420, Figs 41 B – D), Vallone Catrica: 2 valves (AV, BD 421); Sicily: Pezzo: 7 valves (BD 422), Maximum width of the valves: 6.1 / 9.2 / 6.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, moderately elevated, semicarinate, apex well developed, protruding, tail valve semicircular, subcentral protruding mucro. Tegmentum rough, sculptured with numerous oval granules, randomly disposed in HV, LA, PMA, arranged in irregular longitudinal striae in central part of CA, AMA, inclined towards the side margins outside JA, each granule with one subcentral megalaesthete surrounded by 10 – 12 micraesthetes, almost of same size.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	description	Description. Head valve semicircular. Intermediate valves broadly rectangular (W / L = 1.73 – 2.13), moderately elevated (H / W = 0.32 – 0.42), semicarinate in anterior profile, anterior margin convex, side margins rounded, posterior margin practically straight, except for protruding, well developed apex in middle, lateral areas not raised, not differentiated from central area. Tail valve semicircular (W / L = 1.45 – 1.60), anterior margin straight between apophyses, subcentral protruding mucro, antemucronal slope straight or slightly convex, postmucronal slope practically straight, only slightly concave just below mucro. Tegmentum surface rough, space between striae of granules reduced. HV, LA and PMA uniformly sculptured with numerous, oval granules, randomly disposed, (length up to 130 µm), very dense and close to each other. CA and AMA sculptured with oval granules (length up to 137 µm), arranged in irregular longitudinal striae in central part of valve (for about ⅓ of total width), inclined towards the side margins outside JA and much more irregular, granules smaller and denser in JA. Each granule contains one subcentral megalaesthete surrounded by 10 – 12 micraesthetes regularly disposed along edge, megalaesthete slightly larger in some cases, but almost same width. Articulamentum with apophyses rather large, broadly triangular with rounded top, separated by an almost straight jugal sinus, about a third or a little more than valve’s width, connected to primordium of insertion plates in intermediate valves, insertion plates weakly developed in head and tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	discussion	Remarks. The fossil record of Hanleya fratrum sp. nov. is limited to the Pleistocene of South Italy, a few localities from Calabria (Musalà and Vallone Catrica) and Sicily (Pezzo and Salice). Comparisons. This species is at a first check like Hanleya hanleyi (Bean in Thorpe, 1844), from which, however, it is always recognizable for the greater irregularity of the striae of longitudinal granules (slightly larger, up to 137 µm in H. fratrum sp. nov. vs. up to 100 µm in H. hanleyi), for the smaller and denser granules in JA and for the longitudinal striae closest to each other, as opposed to the greater width of the intercostal spaces present in H. hanleyi.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FFB04E700FADFDF86FE990E9.taxon	distribution	Distribution. Pleistocene: central Mediterranean, S. Italy: Musalà, Pezzo, Salice, Vallone Catrica (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8F4E710FADF9B76B3292D3.taxon	description	Fig. 42	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8F4E710FADF9B76B3292D3.taxon	materials_examined	Type material. Holotype SMF 250038, a tail valve Fig. 42 C. Paratypes SMF 250040, a head valve Fig. 42 A and SMF 250039, an intermediate valve Fig. 42 B. Type locality. Höllkopf near Glimmerode (Germany). Type stage. Oligocene (Chattian). Material examined. Lower Oligocene: France: Gaas (Espibos): 6 valves (DA, Figs 42 D, 42 I – L); Gaas (Lagouarde): 1 valve (DA). Oligocene, Chattian: Germany: Kassel, Glimmerode: 3 valves (BD 423, Figs 42 E – H). Maximum width of the valves: 2.2 / 4.7 / 3.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8F4E710FADF9B76B3292D3.taxon	description	Description. Head valve nearly semicircular, highly arched, slightly bent in middle. Intermediate valve broadly rectangular (W / L = 1.72 – 2.28), semicarinate in anterior profile, moderately elevated (H / W = 0.40 – 0.48), anterior margin slightly convex, side margins rounded, posterior margin straight with apex not evident, lateral areas not raised, only marked by a difference in sculpture. Tail valve elliptical, mucro subcentral, blunt, clearly highlighted, antemucronal slope slightly convex, postmucronal slope concave. Tegmentum surface rough. HV, LA and PMA uniformly sculptured with numerous roundish granules, unclearly arranged concentric rows. CA and AMA sculptured with longitudinal striae of roundish to oval granules (ca 10 on each PA, ca 40 in whole CA), straight or slightly obliquely running, finer, more irregular and close set on JA. Articulamentum with apophyses wide, rounded, widely projecting.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8F4E710FADF9B76B3292D3.taxon	discussion	Remarks. The fossil record of Hanleya glimmerodensis Janssen, 1978 is limited to the Oligocene of the Aquitaine Basin (Rupelian) and of the Mainz Basin (Chattian). The only differences highlighted between the valves from Gaas and those from Germany regards the sculpture of tegmentum in JA, well evident in the material from Gaas, more insubstantial or less defined in the material from Germany, and the more elevated intermediate and tail valves from Gaas compared to the German ones (e. g., H / W = 0.43 – 0.48 vs. 0.40). Comparisons. This species is similar to Hanleya hanleyi (Bean in Thorpe, 1844) and H. multigranosa (Reuss, 1860), with which it shares the same type of tegmental sculpture, and from which it differs by slight differences in the valves shape and by the different geographic and stratigraphic range, from extant Mediterranean Sea and Atlantic Ocean up to the upper Miocene of North Italy for H. hanleyi, Middle Miocene of Paratethys for H. multigranosa, and Oligocene of France and Germany for H. glimmerodensis Janssen, 1978.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8F4E710FADF9B76B3292D3.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2018 a); Upper Oligocene: North Europe: Mainz Basin, Germany: Glimmerode, Freden, Söllingen, Doberg (Janssen 1978).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8D4E740FADFF0569C895E8.taxon	description	Fig. 43	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8D4E740FADFF0569C895E8.taxon	description	? (reported as Acanthopleura hanleyi without description or figures); Monterosato 1879, p. 27.? (reported as Hanleya hanleyi without description or figures); Vazzana 1996, p. 148; Long & Zalasiewicz 2011, p. 66, App. 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8D4E740FADFF0569C895E8.taxon	materials_examined	Type material. Syntype, Scarborough Museums Trust, Woodend, Scarborough, U. K. (Sirenko et al. 2016). Type locality. Scarborough (Yorkshire, England). Material examined. Upper Miocene (Tortonian): France: Saint-Clément-de-la-Place: 25 valves (MNHN. F. A 67078, Figs 43 B – D), MNHN. F. A 67079, RGM. 1008399 – 1008400, RGM. 1008430, BD 136). Italy: Borelli: 1 valve (MGPT PU 135038), Rio di Bocca d’Asino: 2 valves (BD 424, MZB 32016). Lower Pliocene: Italy: Liguria: Bussana: 1 valve (BD 425). Pliocene: Portugal: Vale de Freixo: 2 valves (RGM. 1364001, GeoFCUL VFX. 03.364). Upper Pliocene to lower Pleistocene: France: St. Julien-de-Concelles: 1 valve (RGM. 1008441). Pleistocene: Italy: Tuscany: Capraia Island-Capo Corso - 350 m: 3 valves (BD 426); Puglia: SE 06 - 19: 1 valve (MZUB, Fig. 43 E – F); Calabria: Archi S. Francesco: 7 valves (BD 427, Figs 43 A), Musalà: 4 valves (BD 428), Terreti: 1 valve (BD 429), Vallone Catrica: 1 valve (BD 430); Sicily: Ficarazzi: 1 valve (BD 431), Salice: 22 valves (BD 432, Figs 43 G – H). Maximum width of the valves: 2.3 / 5.5 / 4.1 mm. (15.8 / 19 / 18.9 for extant H. nagelfar, see Sirenko 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8D4E740FADFF0569C895E8.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, notched in middle. Intermediate valves broadly rectangular (W / L = 1.87 – 2.10), moderately elevated (H / W = 0.30 – 0.42), carinate in anterior profile, anterior margin undulating, side margins evenly rounded, posterior margin practically straight, except for protruding, well developed apex in middle, lateral areas not raised, only marked by a difference in sculpture. Tail valve somewhat elliptical (W / L = 1.53 – 1.72), anterior margin slightly convex between apophyses, central protruding rounded mucro, antemucronal slope slightly convex, postmucronal slope practically straight, only slightly concave just below mucro. Tegmentum surface slightly rough, space between striae of granules large. HV, LA and PMA uniformly sculptured with numerous, oval granules, randomly disposed, increasing in size and spacing towards outer margins. CA and AMA sculptured with roundish to oval granules (length 75 – 100 µm), arranged in longitudinal striae, fine and close set on JA, getting larger and posteriorly converging towards side margins. Each granule contains one central megalaesthete surrounded by 8 – 12 micraesthetes, megalaesthete slightly larger in some cases, but almost same width. Articulamentum with apophyses rather large, broadly triangular with rounded top, separated by a slightly concave jugal sinus, about ⅓ of valve ‘ s width, connected to primordium of insertion plates in intermediate valves, insertion plates weakly developed in head and tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8D4E740FADFF0569C895E8.taxon	discussion	Remarks. This species has a complex taxonomic history, and many synonyms are known and accepted (Sirenko et al. 2016). Several publications reported Hanleya hanleyi Bean in Thorpe, 1844 as occurring in the Mediterranean Sea and in deposits from Neogene of Italy (Sabelli 1972, 1974 a; Dell’Angelo & Smriglio 1999; Dell’Angelo et al. 2015 a, 2021 a). Recently, another species of Hanleya (H. mediterranea Sirenko, 2014) has been described from the Mediterranean Sea (Dell’Angelo et al. 2021 a), and both species occur in fossil deposits from Italy, at places together (e. g., in the Upper Miocene of Borelli, see Dell’Angelo et al. 2015 a). Therefore, for some bibliographical references of fossil H. hanleyi (without description and / or figures) a specific determination (H. hanleyi or H. mediterranea) cannot be reached. Also, a list of bibliographical references for living Hanleya in the Mediterranean Sea has been given by Dell’Angelo et al. (2021 a: 133), under the taxon “ Hanleya sp. indet. ”. The sculpture shows a certain variability, as has been reported by Dell’Angelo et al. (2021 a) on recent species, the longitudinal series of granules are more or less regular, some split (Dell’Angelo et al. 2021 a: fig. 1. D) and some converge towards the jugal area (Figs 43 B, 43 G), the interstices can be more or less narrow.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8D4E740FADFF0569C895E8.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b); Proto – Mediterranean Sea (Tortonian): N. Italy: Borelli, Rio di Bocca d’Asino (Dell’Angelo et al. 1999, 2015 a). Lower Pliocene: North Europe, Belgium: Kallo (Marquet 1984, 2002); northeastern Atlantic: U. K.: Sutton (Wood 1848; Reid 1890); central Mediterranean, Italy: Liguria: Bussana (this study). Pliocene: northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo et al. 2022). Upper Pliocene to lower Pleistocene: northeastern Atlantic (Zanclean): Anjou, France: St. Julien-de-Concelles (Dell’Angelo et al. 2018 b). Pleistocene: central Mediterranean, Italy: Archi S. Francesco, Ficarazzi, Vallone Catrica, Musalà, Salice, Terreti (this study), Capraia Island-Capo Corso (Dell’Angelo & Giusti 1997), SE 06 (Taviani et al. 2023). Recent: North and Central Atlantic Ocean, near southern Greenland (Hansson 1998; Sneli & Gudmundsson 2018), North America, Europe (Kaas 1979; McKay & Smith 1979; Consolado Macedo et al. 1999; Urgorri et al. 2017), Canary Islands (Kaas 1991; Hernández & Rolán 2011) and northern Africa, Madeira Arch. (Kaas 1991; Segers et al. 2009); Mediterranean Sea: Italy (Dell’Angelo et al. 2021 a), Aegean Sea (Zenetos & Van Aartsen 1995), Turkey (Ozturk et al. 2014) (Sirenko 2014; Sirenko et al. 2016; Dell’Angelo et al. 2021 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8B4E750FADFEB46B5F912C.taxon	description	Fig. 44	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8B4E750FADFEB46B5F912C.taxon	materials_examined	Type material. Holotype ZISP 2201, length 4.7 mm, now disarticulated, and three paratypes ZISP 2202. Type locality. Mediterranean Sea, off Begur (Girona, Spain), 200 – 300 m. Material examined. Upper Miocene (Tortonian): Italy: Borelli: 1 valve (MGPT PU 135039); Montegibbio: 2 valves (BD 433, MZB 32017, Figs 44 G – H). Lower Pliocene: Italy: Borzoli: 4 valves (BD 434), Rio Sant’ Antonino: 3 valves (MP, MZB 45706), Zinola: 1 valve (MZB 45705). Pliocene: Spain: Estepona: 1 valve. Italy: Sicily: Altavilla: 1 valve (BD 435, Figs 44 E – F), Trappeto: 1 valve (BD 436). Pleistocene: Italy: Tuscany: Capraia Island-Capo Corso - 350 m: 4 valves (BD 437, Figs 44 A – C), Riparbella: 2 valves (BD 438, Fig. 44 D); Calabria: Archi S. Francesco: 1 valve (BD 439). Maximum width of the valves: 3 / 3.8 / 2.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8B4E750FADFEB46B5F912C.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, notched in middle. Intermediate valves trapezoidal, width about twice length (W / L = 1.88 – 1.94), moderately elevated (H / W = 0.40 – 0.45), semicarinate in anterior profile, anterior margin greatly splayed laterally, with large inward curve in jugal area, posterior margin with protruding, well developed apex in middle, lateral areas not raised, only marked by a difference in sculpture, Tail valve more or less elliptical (W / L = 1.43 – 1.58), anterior margin slightly convex between apophyses, mucro subcentral, clearly indicated, antemucronal slope slightly convex, postmucronal slope decidedly concave directly behind mucro, straightening near posterior margin. Tegmentum surface smooth, without aesthetes among granules. HV, LA and PMA uniformly sculptured with roundish granules (diameter 70 – 80 μm), arranged without pattern. CA and AMA sculptured with oval granules (120 × 70 μm) arranged without pattern, several granules join to form larger granules (up to 220 × 160 μm), JA sculptured with oval granules (about 80 × 50 μm) arranged without interspaces or pattern. Each granule contains one central megalaesthete surrounded by 9 – 16 micraesthetes, megalaesthete slightly larger in some cases, but almost same width. Articulamentum strongly developed, apophyses wide, oriented laterally, connected to primordium of insertion plates in intermediate valves, insertion plates obsoletely striate in head and tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8B4E750FADFEB46B5F912C.taxon	discussion	Remarks. The fossil record of Hanleya mediterranea Sirenko, 2014 includes the Miocene (Tortonian) of N. Italy, Pliocene of Italy and Spain, and Pleistocene of Italy. Hanleya mediterranea has been recently described from the Mediterranean Sea, and both the species living in the Mediterranean Sea [H. mediterranea and H. hanleyi (Bean in Thorpe, 1844)] has been found in fossil deposits from Italy, in some case with both species present in the same site (e. g., in the Upper Miocene of N. Italy, see Dell’Angelo et al. 2015 a). Comparisons. Hanleya mediterranea is distinguished from other species in the genus by the presence of large granules comprising two or more small granules in pleural areas. In H. mediterranea the head valve has more strongly developed insertion plates that are 16 – 20 % the length of the valve, compared with H. hanleyi in which the insertion plates are 3 – 9 % the length of the head valve (Sirenko 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8B4E750FADFEB46B5F912C.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, North Italy: Borelli, Montegibbio (Dell’Angelo et al., 1999, 2015 a). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Rio S. Antonino, Zinola (Dell’Angelo et al. 2013). Pliocene: western Mediterranean, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Altavilla Milicia (Dell’Angelo et al. 2012). Pleistocene: central Mediterranean, Italy: Riparbella (Dell’Angelo & Forli 1995 a), Torrente Stirone (Sabelli & Taviani 1979), Capraia Island-Capo Corso - 350 m, Archi S. Francesco (this study). Recent: Mediterranean Sea (Sirenko 2014; Sirenko et al. 2016; Dell’Angelo et al. 2021 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8A4E770FADFBF068E89604.taxon	description	Fig. 45	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8A4E770FADFBF068E89604.taxon	materials_examined	Type material. Unknown, not present in NHMV. Type locality. Rudoltice (Czech Republic). Type stage. Middle Miocene. Material examined. Middle Miocene: Central Paratethys: Czech Republic: Porzteich: 1 valve (NHMV 2010 / 0256 / 0014, Figs 45 E – H). Maximum width of the valves: 2.6 / 4.2 / 2.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8A4E770FADFBF068E89604.taxon	description	Description. Head valve semicircular, quite narrow and strongly arched. Intermediate valves broadly rectangular, semicarinate in anterior profile, side margins rounded, posterior margin almost straight, except for well developed apex in middle, lateral areas not raised. Tail valve elliptical, mucro subcentral, clearly indicated. Tegmentum surface smooth, deprived of aesthetes among granules. HV, LA and PMA uniformly sculptured with numerous roundish granules, more irregularly arranged and more variable in shape in LA. PA and AMA sculptured with elliptical granules (up to 170 x 70 µm), arranged in longitudinal very irregular striae, granules smaller (up to 130 x 50 µm) and more closely spaced in JA, transition from small to larger granules not gradual, but rather abrupt. Each granule contains 1 megalaesthete and up to 25 microaesthetes or more, megalaesthete slightly larger in some cases, but almost same width, no aesthetes among granules. Articulamentum with apophyses wide, rounded, insertion plates well developed both on head and tail valves, slightly pectinated on head valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8A4E770FADFBF068E89604.taxon	discussion	Remarks. Hanleya multigranosa (Reuss 1860) is a rare and poorly known species. Reuss (1860) described it on the basis of two valves from Rudoltice, a head valve (Fig. 45 A) and a tail one, this last (Reuss 1860: pl. 8, figs 9 a – b) with a sculpture not matching any of the species known from the Oligocene to Pleistocene of Europe, and probably not attributable to the genus Hanleya (see the discussion in Šulc). Šulc (1934) described a few valves (head, intermediate and tail, Figs 45 B – D) from Rudoltice, indicating also two other sites for this species, Kostej (Romania) and Porzteich (Czech Republic), and noted that the tail valve illustrated by Reuss differs considerably from the tail valves attributed by him to Hanleya multigranosa. De Rochebrune (1882) and Procházka (1900) do not figure this species but refers to the illustrations of Šulc (1934). Bałuk (1971, 1984) described many valves from Korytnica (Poland), attributed to H. multigranosa following the Šulc’s descriptions and illustrations, also regarding the tail valve, and therefore not the one figured by Reuss (1860: pl. 8, figs 9 a – 9 b). The last known record of H. multigranosa is that of Ruman & Hudácková (2015), a few valves from Rohožník (Slovakia). A single incomplete intermediate valve of Hanleya multigranosa from Porzteich (Czech Republic), is preserved at NHMW in the Šulc collection (Figs 45 E – H), but not registered as the type material. The sculpture of the tegmentum is consistent with the attribution to the genus Hanleya, also if the articulamentum’s features are not visible: the granules are elevated, well separated, with a high number of micraesthetes around a megalaesthetes, arranged in irregular rows longitudinally oriented in the median area, and irregularly covering the jugal area. The sculpture of this valve corresponds to that described by Šulc (1934), apart for the shape of the granules. Šulc’s description says “ Some (granules, on the median valve) are elongated in longitudinal direction, but they usually have a circular cross section ”. The granules in the examined valve are all of elliptical form, but in Hanleya spp. (e. g., H. hanleyi) the granules in intermediate valves tend to be more elongated than the roundish granules predominant in HV and PA (see Dell’Angelo et al. 1998 a: pl. 2, figs 1 – 2 and Dell’Angelo & Smriglio 1999: pl. 25, figs A, C, E, pl. 26, figs I, J). The Šulc’s figures of intermediate valve are not so clear to resolve the doubts, and also the description of intermediate valves by Ashby & Cotton (1935) does not give any supplementary information. Notwithstanding, there are no other species of Hanleya described for the Miocene of Paratethys, and the locality (“ Porzteich ”) is one of those reported by Šulc, so we attribute the examined valve to H. multigranosa. Some authors consider Hanleya multigranosa to be the ancestral species of the Recent species Hanleya hanleyi (Bean in Thorpe, 1844) (Šulc 1934; Malatesta 1962; Laghi 1977). In fact, the two species are rather similar). Unfortunately we were unsuccessful in locating the type material of H. multigranosa, which is not in the NHMW, the description given by Reuss (1860) is incomplete, and also the subsequent integration by Šulc (1934), as important diagnostic information is missing and his tail valve do not correspond to that described by Reuss, and for these reasons, and the conspicuous difference in stratigraphic and geographic distribution, we consider H. multigranosa and H. hanleyi as distinct species. The elliptical granules of the valve figured from Porzteich (Figs 45 E – H) are variable in length, up to 170 μm, while the granules of intermediate valves of H. hanleyi range up to 100 μm (Sirenko et al. 2016: figs 2 E, 5 E). Also, the number of micraesthetes around the megalaesthete is different, 8 – 12 for H. hanleyi (Dell’Angelo & Smriglio 1999: 86; Sirenko et al. 2016: figs 2 E, 5 E) vs. up to 25 or more in the valve of H. multigranosa examined (but up to 14 in the valves from Korytnica studied by Bałuk 1984). The longitudinal rows of granules in the pleural areas of intermediate valves of H. hanleyi are more regular than in the valve examined, the granules are closer to one another and give a different look to the sculpture of the valve (see Sirenko et al. 2016: figs 2 B, 5 B – C). Comparisons. Hanleya multigranosa differs from H. mediterranea Sirenko, 2014 in having a shorter insertion plate on the head valve, a differently shaped tail valve, and smaller and more numerous granules with a larger number of micraesthetes (up to 25 or more vs. 12 – 13 in H. mediterranea).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF8A4E770FADFBF068E89604.taxon	distribution	Distribution. Lower-Middle Miocene: North Europe: Belgium: Antwerp (Louwye et al. 2010). Middle Miocene: Central Paratethys (Langhian-Serravallian): Czech Republic: Rudoltice, Porzteich (Reuss 1860; Procházka, 1900; Šulc 1934; this study), Slovakia: Rohožník (Ruman & Hudácková 2015), Romania: Coştei (Šulc 1934), Poland: Korytnica (Bałuk 1971, 1984).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF884E780FADFC1869ED933D.taxon	description	Fig. 46	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF884E780FADFC1869ED933D.taxon	materials_examined	Type material. Holotype MNHN. F. A 67080, intermediate valve, width 9 mm, Figs 46 B – D. Paratypes: MNHN. F. A 67081 – A 67086 (6 valves), Figs 46 A, 46 E – H; NHMW 2017 / 0108 / 0012 – 0014 (3 valves); RGM. 1008401 – 1008402 (2 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): France: Saint-Clément-de-la-Place: type material plus 74 valves (MNHN. F. A 67087, NHMW 2017 / 0108 / 0015, RGM. 1008350, RGM. 1008403, BD 137); Sceaux d’Anjou, La Presselière: 2 valves (BD 138). Maximum width of the valves: 7 / 9 / 6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF884E780FADFC1869ED933D.taxon	description	Description. Head valve subovate, posterior margin widely V-shaped. Intermediate valves polygonal (W / L = 1.77 – 1.84), carinate in anterior profile, moderately elevated (H / W = 0.41), posterior margin with small projecting apex, both sides slightly concave, lateral areas not raised. Tail valve elliptical, anterior margin convex, posterior margin rounded, mucro central, strongly elevated, prominent, antemucronal slope slightly convex, postmucronal slope decidedly concave directly behind mucro, straightening near posterior margin Tegmentum surface slightly rough, without aesthetes among granules. HV, LA and PMA uniformly sculptured with scattered roundish granules, elevated, diameter 75 μm. PA and AMA with conspicuous radial depression, sculptured with elongate oval, elevated granules arranged in striae, obliquely irregularly directed, thickening and more longitudinal towards JA, where they become strongly thickened; granules irregular, oval to elongate, oblique, up to 100 μm long in PA, smaller, up to 75 μm and densely packed in JA, up to 160 μm long in AMA obliquely directed. Each granule contains one or two megalaesthetes in central position, and up to 15 or more micraesthetes irregularly disposed in CA and AMA, up to 10 – 12 micraesthetes arranged irregularly along margin in HV, LA and PMA; all aesthetes of same size, no aesthetes among granules. Articulamentum solid, apophyses wide, long, connected to primordium of insertion plates in intermediate valves, insertion plates well developed both on head and tail valves, slightly pectinated on tail valve, muscle scars particularly prominent on tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF884E780FADFC1869ED933D.taxon	discussion	Remarks. The fossil record of Hanleya sancticlementensis Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is limited to the Miocene (Tortonian) of the northwestern France. The material from Saint-Clément-de-la-Place consists mostly of incomplete and poorly preserved valves, especially the characters of the articulamentum are sometimes scarcely visible. Hanleya sancticlementensis is readily distinguished from its congeners by its peculiar sculpture on PA, with irregular striae of granules obliquely directed near LA, becoming thicker and more longitudinal towards JA, with an area where the orientation of the rows is reversed, from oriented slightly to the left to oriented slightly towards the right; the rows of granules almost coalesce on JA. The triangular jugal area has a different sculpture, with rows of more appressed smaller granules. The shape and size of the granules varies on different areas of the valves, as do the number and position of aesthetes. Comparisons. Hanleya sancticlementensis is superficially similar to H. harasewychi Sirenko, 2014, a living species from the eastern Atlantic frontage; it differs from H. harasewychi by the shape of the intermediate valves (W / L = 1.77 – 1.84 vs. 2.3 – 2.7 in H. harasewychi), the sculpture in PA, with granules arranged in longitudinal rows and the lack of pattern in JA of H. harasewychi.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF884E780FADFC1869ED933D.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place, Sceaux d’Anjou, La Presselière (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF874E7A0FADF9C3696C95E8.taxon	description	Fig. 47	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF874E7A0FADF9C3696C95E8.taxon	materials_examined	Type material. Holotype: MZUB 60338, intermediate valve, width 3.8 mm (Figs 47 A – D); Paratype 1: MNHN-IM- 2022 - 2410, intermediate valve, width 3.4 mm from offshore Capraia Island-Capo Corso - 350 / 500 m; Paratype 2: MZUB 60339, intermediate valve from Tipaza, Algeria, width 3.5 mm; Paratype 3: SMF 66407, intermediate valve from Gorgona island, width 3.5 mm; Paratype 4: ZISP 2423, tail valve, width 2.7 mm from Scilla. Type locality. Tyrrhenian Sea, Scilla (Calabria, Italy). Material examined. Pleistocene: Italy: Capraia Island-Capo Corso - 350 / 500 m: type material, Archi S. Francesco: 1 valve (BD 440, Figs 47 E – H). Recent: Italy: type material, plus Tuscan Archipelago: Capraia Island: 3 valves (BD 249, ZISP 2424), Giglio Island: 1 valve (BD 250); Algeria: Tipaza: type material. Maximum width of the valves: -- / 4.0 / 3.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF874E7A0FADF9C3696C95E8.taxon	description	Description. Head valve unknown. Intermediate valves trapezoidal (W / L = 1.50 – 1.84), elevated (H / W = 0.39 – 0.53), semicarinate in anterior profile, anterior margin greatly splayed laterally, almost straight to slightly concave in jugal area, posterior margin with protruding, well developed apex in middle, lateral areas not raised, only marked by a difference in sculpture, Tail valve circular (W / L = 1.16 – 1.32), anterior margin almost straight between apophyses, mucro submedian, antemucronal slope convex, postmucronal slope decidedly concave directly behind mucro, straightening near posterior margin. Tegmentum surface slightly rough, without aesthetes among granules. LA and PMA uniformly sculptured with roundish granules (diameter up to 100 μm), arranged without pattern. PA and AMA sculptured with oval granules (up to 127 μm long) arranged without pattern, JA sculptured with smaller granules, more irregular and closer to each other, without interspaces. Each granule contains one megaesthete in central position surrounded by up to 15 others in CA, AMA, a little less (up to 12) in LA, PMA; all aesthetes more or less of same size, no aesthetes among granules. Articulamentum strongly developed, apophyses wide, oriented laterally, connected to primordium of insertion plates in intermediate valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF874E7A0FADF9C3696C95E8.taxon	discussion	Remarks. Hanleya schwabei Dell’Angelo, Sosso & Taviani, 2024 has been recently established upon a few valves from various Mediterranean localities, previously labeled as H. aff. mediterranea Sirenko, 2014 in collections. Some such valves were illustrated by Dell’Angelo et al. (2021: figs 5. A – F). A single intermediate valve offshore CapraiaIsland-Capo Corso - 350 / 500 m could belong to last glacial Pleistocene assemblages. We found some additional loose valves from the Tuscan archipelago, and a tail valve from the Pleistocene of Archi S. Francesco (Figs 47 E – H), confirming the presence of H. schwabei as fossil in the Italian Pleistocene. Comparisons. The most similar species is H. mediterranea Sirenko, 2014, that shares the tegmentum sculpture characterized by granules arranged without pattern, but differ anyway from H. schwabei by the different sculpture of the tegmentum, with characteristic large granules not present in H. schwabei, which give a different appearance to the valves. Moreover the shape of the tail valves is different, almost circular in H. schwabei, more or less elliptical in H. mediterranea.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF874E7A0FADF9C3696C95E8.taxon	distribution	Distribution. Pleistocene: central Mediterranean, Italy: Archi S. Francesco (this study), Capraia Island-Capo Corso - 350 / 500 m (Dell’Angelo et al. 2024). Recent: Mediterranean Sea: Italy: Capraia Island, Giglio Island, Scilla and Algeria: Tipaza (Dell’Angelo et al. 2024; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	description	Fig. 48	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	materials_examined	Type material. Holotype MSNG 62635, intermediate valve, width 11.5 (14.4) mm (Figs 48 A – H). Type locality. Vallone Catrica, Reggio Calabria (Calabria, Italy). Type stage. Lower Pleistocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	etymology	Etymology. The specific name honors Boris Sirenko (Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia), for his prominent contribution to the study of Recent and fossil chitons.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	materials_examined	Material examined. Pleistocene: Italy: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	diagnosis	Diagnosis. Intermediate valves broadly rectangular, moderately elevated, carinate, small protruding apex. Tegmentum surface barely visible, granules very close to each other, arranged without pattern, more roundish and a little smaller in LA, oval in CA, each granule with 1 – 3 megaesthete in central position surrounded by up to 20 – 25 microaesthetes, all of same size. Articulamentum solid, apophyses wide.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	description	Description. Head and tail valve unknown. Intermediate valves broadly rectangular, moderately elevated (H / W estimated = 0.38), carinate in anterior profile, side margins rounded, posterior margin almost straight with protruding, small apex in middle, lateral areas not raised, Tegmentum surface barely visible, granules very close to each other. LA uniformly sculptured with more roundish and slightly smaller granules, arranged without pattern. CA sculptured with oval granules (180 – 210 μm long) arranged without pattern and very close, granules of similar shape and size on PA and JA. Each granule contains 1 – 3 megaesthete in central position surrounded by up to 20 – 25 microaesthetes in CA, little less (up to 15) in LA, all aesthetes of same size. Articulamentum solid, apophyses wide, long, connected to primordium of insertion plates in intermediate valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	discussion	Remarks. The fossil record of Hanleya sirenkoi sp. nov. is limited to the Pleistocene of S. Italy. Our material is represented by a unique intermediate valve not complete, but with a series of characteristics very different from the other spp. of Hanleya described here and such as to justify the establishment of a new species. Notwithstanding the incompletness of the unique valve, we have estimated the total width (put in brackets after the measured value) and the dorsal elevation H / W of the complete valve. In some granules the size of the pores of the megalaesthetes seem slightly larger than those of microaesthetes, but this could be due to erosion, for which we rate megalaesthetes and microaesthetes roughly of the same size. Comparisons. Hanleya sirenkoi sp. nov. shares the sculpture consisting of granules irregularly arranged without pattern with H. mediterranea Sirenko, 2014, H. schwabei Dell’Angelo, Sosso & Taviani, 2024 and H. sossoi Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018, from which it differs for the larger and very close granules (only H. sossoi has granules of similar size, but with a different arrangement) and the distinct structure of the aesthetes.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF854E7B0FADFEB46EEE94C8.taxon	distribution	Distribution. Pleistocene: central Mediterranean, S. Italy: Vallone Catrica (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF844E7C0FADFDD46E5191D9.taxon	description	Fig. 49	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF844E7C0FADFDD46E5191D9.taxon	materials_examined	Type material. Holotype MNHN. F. A 67088, intermediate valve, width 4 mm (Figs 49 A – D). Paratypes: MNHN. F. A 67089 – A 67091 (3 valves, Figs 49 E – G); NHMW 2017 / 0108 / 0016 – 0018 (3 valves); RGM. 1008404 – 1008405 (2 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): France: Saint-Clément-de-la-Place: type material plus 110 valves (MNHN. F. A 67092, Fig. 49 H, MNHN. F. A 67093, NHMW 2017 / 0108 / 0019, RGM. 1008351, RGM. 1008406, RGM. 1008431, BD 139), Sceaux d’Anjou, La Presselière: 1 valve (RGM. 1008446), Beugnon: 1 valve (RGM. 1310178). Maximum width of the valves: 6.1 / 6 / 7.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF844E7C0FADFDD46E5191D9.taxon	description	Description. Head valve subovate, posterior margin widely V-shaped. Intermediate valves polygonal, width about twice length (W / L = 1.92 – 2.03), carinate in anterior profile, moderately elevated (H / W = 0.38), posterior margin with small projecting apex, both sides slightly concave, lateral areas not elevated. Tail valve from elliptical to circular (W / L = 1.31 – 1.47), anterior margin convex, posterior margin rounded, mucro central, strongly elevated, prominent, antemucronal slope slightly convex, postmucronal slope decidedly concave directly behind mucro, straightening near posterior margin. Tegmentum surface smooth, granules more spaced from each other, there are no aesthetes among granules. HV, LA and PMA uniformly sculptured with roundish to elongate granules, arranged without pattern, roundish in HV (diameter ca. 100 μm towards margin, progressively smaller towards apex), roundish to slightly elongate in PMA (up to 140 μm long), elongate in LA (up to 130 μm long). CA and AMA with slight radial depression, sculptured with elongate, elevated granules irregularly arranged, but giving the impression of a very irregular longitudinal arrangement, more oblique towards lateral areas, up to 175 μm long in CA, up to 220 μm long in AMA. Variable number of megalaesthetes and microaesthetes, all of same size, up to 30 – 35 irregularly disposed in CA, 3 – 5 megalaesthetes in central position and up to 20 – 25 microaesthetes or more irregularly disposed along margin in AMA, 20 or more irregularly distributed in HV, 10 – 15 in LA, 2 – 4 megalaesthetes in central position and up to 12 microasthetes irregularly disposed along margin in PMA; all aesthetes of same size, no aesthetes among granules. Articulamentum solid, apophyses wide, long, connected to primordium of insertion plates in intermediate valves, insertion plates well developed both on head and tail valves, pectinated on tail valve, muscle scars particularly prominent on tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF844E7C0FADFDD46E5191D9.taxon	discussion	Remarks. The fossil record of Hanleya sossoi Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is limited to the Miocene (Tortonian) of the northwestern France. The material from the Miocene (Tortonian) of Saint-Clément-de-la-Place consists mostly of incomplete and poorly preserved valves, especially the characters of the articulamentum are sometimes scarcely visible. The shape and size of the granules is variable on the different areas of the valves, but are similar in PA and JA of intermediate and tail valves. The number and position of microaesthetes is also variable. Comparisons. Hanleya sossoi can be compared with the other Hanleya spp. with a tegmental sculpture characterized by granules irregularly arranged without pattern (H. mediterranea Sirenko, 2014, H. schwabei Dell’Angelo, Sosso & Taviani, 2024, and H. sirenkoi sp. nov.). Hanleya sossoi is the species that has larger (up to 220 μm long) and more spaced from each other granules, and also the greatest number of aesthetes (up to 30 – 35).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF844E7C0FADFDD46E5191D9.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Beugnon, Saint-Clément-de-la-Place, Sceaux d’Anjou, La Presselière (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF834E7D0FADFAEC6E04947C.taxon	type_taxon	Type species. Chiton laevis Montagu, 1803 (non Pennant, 1777) = Callochiton septemvalvis (Montagu, 1803), by subsequent designation (Gray 1847)	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF834E7D0FADFAEC6E04947C.taxon	distribution	Distribution. Callochiton is known from the Oligocene up to the Recent, with a living distribution in the Indo-West Pacific (including Japan), eastern Atlantic Ocean, and subantarctic waters (Kaas & Van Belle 1985 b). The fossil record includes the Oligocene of New Zealand (Ashby 1929; Lee et al. 2014; Wu & Lee 2024), the Miocene of Europe (Šulc 1934; Bałuk 1984; Dell’Angelo et al. 2016), Argentina (Urteaga et al. 2011) and Australia (Ashby 1939), the Pliocene / Pleistocene of Europe (Koskeridou et al. 2009; Dell’Angelo et al. 2011, 2004, 2013), New Zealand (Sutherland et al. 1995), the Pleistocene of Japan (Itoigawa et al. 1976), Red Sea (Dell’Angelo et al. 2020 a), Greece (Garilli et al. 2005), and Italy (Sabelli & Taviani 1979; Dell’Angelo & Giusti 1997), and the Holocene of Patagonia (Gordillo & Schwabe 2009).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF834E7D0FADFAEC6E04947C.taxon	discussion	Remarks: Recent molecular phylogenetic analyses support the division of Polyplacophora into three orders, Lepidopleurida, Callochitonida and Chitonida (Giribet & Edgecombe, 2020; Irisarri et al. 2020; Moles et al. 2021; Liu, Sigwart & Sun 2023). Three extant species of Callochiton are known from the Neogene and Pleistocene of the circum-Mediterranean area: C. septemvalvis (Montagu, 1803), C. doriae (Capellini, 1859), and C. calcatus Dell’Angelo & Palazzi, 1994, and other two species only known as fossils are described from the Neogene of Europe: Callochiton pouweri Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 and C. zigzag Šulc, 1934. The main morphological characters of the Callochiton spp. considered in the present study are reported in Tab. 8.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF814E7F0FADFF0569BD912C.taxon	description	Fig. 50	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF814E7F0FADFF0569BD912C.taxon	materials_examined	Type material. Holotype MZB 11620, one specimen (length 4.5 mm). Paratypes: Dipartimento di Scienze della Terra of Milan University, reg. n. 7040 – 7044, 5 spm from Isole Pontine); private collections: AL (1 spm from Favignana Island), BD 4283 (1 spm from Isole Pontine), BD P 50 / 4 (10 valves from Marettimo Island), BD P 152 / 3 (2 valves from Northern Tyrrenian Sea), BD P 157 / 2 (1 valve from Elba Island), EM (1 spm from Capo Passero), LB (1 spm from Trapani), LD A 199 (1 spm from Mallorca Island), LD A 208 (3 spm from Mallorca Island), A 213 (2 spm from Mallorca Island), LD A 256 (2 spm from Mallorca Island), LD A 412 (1 spm from Formentera). Type locality. Villasimius (Cagliari) ex nets at 80 – 100 m. Material examined. Pliocene: Italy: Tuscany: Poggio alla Fame: 10 valves (BD 573, Figs 50 A – G). Pleistocene: Italy: Tuscany: Capraia Island-Capo Corso - 350 / 500 m: 62 valves (BD 574, Figs 50 H – I); Calabria: Petti di Carrubbare: 8 valves (BD 575, Figs 50 J – L); Sicily: Capo Milazzo: 2 valves (BD 576), Casa Catarinicchia: 5 valves (BD 577), Casa Parrino: 6 valves (BD 578). Recent: Italy: type material from Elba Island (Grosseto), Isole Pontine (Latina), Villasimius (Cagliari), Capo Passero (Siracusa), Favignana Island (Trapani), Trapani; Spain: type material from Formentera and Mallorca Island. Maximum width of the valves: 2.5 / 2.5 / 1.9 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF814E7F0FADFF0569BD912C.taxon	description	Description: Head valve semicircular, front slope convex. Intermediate valves broadly rectangular (W / L = 1.51 – 1.98), semicarinate in anterior profile, decidedly elevated (H / W = 0.50 – 0.75), anterior margin convex to sinuose, side margins rounded, posterior margin concave at both sides of strongly protruding, obtuse apex, lateral areas raised, quite evident. Tail valve semicircular / elliptical (W / L = 1.44 – 1.50), anterior margin convex, mucro in anterior position, not prominent, antemucronal slope almost straight, postmucronal slope slightly convex. Tegmentum granulose, rough. HV, LA and PMA sculptured with fine granules, fuse into continuous lines radially, and with characteristic black dots, pigmented cusps of shell-eyes, irregularly distributed but with prevalence towards margins. CA and AMA sculptured with longitudinal lines of micraesthetes, with 6 – 8 conspicuous longitudinal grooves in front of diagonal ridge, grooves shortening towards jugum Articulamentum with apophyses wide, short, regularly rounded, connected at jugum by a relatively long jugal plate, slit formula 15 – 16 / 2 / 12 – 13, teeth irregular, slit rays evident, with large pores, eaves coarsely porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF814E7F0FADFF0569BD912C.taxon	discussion	Remarks. The fossil record of Callochiton calcatus Dell’Angelo & Palazzi, 1994 is limited to the Pliocene and Pleistocene of Italy. This extant species is easily distinguished by the mucro in anterior position, the postmucronal slope convex noticeable dorsal and the noticeable elevation of its intermediate valves. We have attributed to Callochiton calcatus some valves from the Pliocene of Poggio alla Fame showing a slightly lower H / W ratio (0.50 – 0.56, Figs 50 C – D, vs. 0.65 – 0.72 reported in the original description of the species). The other characters are those typical of C. calcatus, e. g., the mucro’s position of the tail valve and the convexity of the postmucronal slope (Figs 50 G), and so we can consider the H / W ratio to be highly variable, in a range from 0.50 (Fig. 50 C) to 0.75 (Fig. 50 I). Comparisons. Differences with Callochiton pouweri are discussed below.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF814E7F0FADFF0569BD912C.taxon	distribution	Distribution. Pliocene: central Mediterranean, Italy: Poggio alla Fame (Chirli 2004; this study). Pleistocene. central Mediterranean, Italy: Capraia Island-Capo Corso - 350 / 500 m (Dell’Angelo et al. 2001 a), Capo Milazzo, Casa Catarinicchia, Casa Parrino, Petti di Carrubbare (this study). Recent. Mediterranean: random records from the Balearic Islands (Mallorca, Formentera) to the Tyrrhenian coasts of Italy, Sardinia and Sicily (Dell’Angelo & Smriglio 1999), Strait of Sicily: Graham Bank (Dell’Angelo et al. 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF804E620FADFBF069A19108.taxon	description	Fig. 51 Chiton euplaeae O. G. Costa, 1830, p. i, iv, pl. 1, fig. 3 (nomen dubium).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF804E620FADFBF069A19108.taxon	materials_examined	Type material. Unknown. Not present at MSNG (M. Tavano, pers. com.). Type locality. Gulf of La Spezia. Material examined. Lower Miocene: Italy: Sciolze (Burdigalian): 3 valves (MZB 32059, PG), Fig. 51 E. Middle Miocene: Italy: Albugnano (Langhian): 2 valves (MZB 32060, Figs 51 F – H, PG); Central Paratethys (Langhian-Serravallian): Hungary: Bánd: 1 valve (BD 579, Letkés: 1 valve (BD 580); Romania: Lapugy: 1 valve (BD 581); Eastern Paratethys: Ukraine: Varovtsi: 3 valves (BD 582, Figs 51 A – B). Upper Miocene (Tortonian): France: Saint-Clément-de-la-Place: 162 valves (MNHN. F. A 67108 – 110, Figs 51 J – L, NHMW 2017 / 0108 / 0028, RGM. 1008408 – 1008410, RGM. 1008434, BD 145), Renauleau: 9 valves (MNHN. F. A 67111, BD 146). Italy: Rio di Bocca d’Asino: 20 valves (BD 583, MZB 32062 – 32063, PG), Villa Monti: 1 valve (MZB 32061). Lower Pliocene: Italy: Liguria: Borzoli: 63 valves (BD 584), Bussana: 53 valves (BD 585), Caranchi: 8 valves (BD 586, MP, MZB 45726, Fig. 51 I), Garlenda: 1 valve (MP), Genova Sestri: 11 valves (BD 587), Rio Sant’Antonino: 52 valves (MP), Rio Torsero: 5 valves (BD 588, MP, MZB 45725), Zinola: 1 valve (BD 589). Pliocene: Spain: Estepona: 11 valves (BD 590). Portugal: Vale de Freixo: 213 valves (GeoFCUL VFX. 03.340 – 341, GeoFCUL VFX. 03.356 – 357, RGM. 1364012 – 1364013, MNHN. F. A 81987, BD 242). Italy: Emilia-Romagna: Cava di Campore: 11 valves (BD 591), Gagliardella “ Tagliata ”: 2 valves (BD 592), Marano sul Panaro: 2 valves (BD 593); Tuscany: Bibbiano: 3 valves (BD 594); Castell’Anselmo: 4 valves (BD 595); Castiglioncello del Trinoro: 2 valves (BD 596); Colle Val d’Elsa: 4 valves (BD 597); Orciano Pisano: 19 valves (BD 598); Pietrafitta: 51 valves (BD 599); Serre di Rapolano: 2 valves (BD 600); Umbria: Orvieto: 1 valve (BD 601); Sicily: Altavilla: 5 valves (AG, AR, BD 602), Trappeto: 10 valves (BD 603). Upper Pliocene to upper Pleistocene: France: Bosq d’Aubigny: 2 valves (RGM. 1310185). Pleistocene: Greece: Kyllini: 16 valves (BD 604, DGUP, Figs 51 C – D); Italy: Tuscany: Capraia Island-Capo Corso - 350 / 500 m: 27 valves (BD 605), Cisternino: 10 valves (BD 606), Fauglia: 7 valves (BD 607); Riparbella: 75 valves (BD 608); Emilia-Romagna: Torrente Guerro: 1 valve (BD 609), Torrente Stirone: 24 valves (BD 610); Puglia: Cutrofiano: 2 valves (BD 611), Gallipoli: 5 valves (BD 612); Calabria: Archi S. Francesco: 13 valves (BD 613), Carrabbati: 2 valves (BD 614), Gallina: 8 valves (BD 615), Le Castella: 33 valves (BD 616), Musalà: 2 valves (BD 617), Pecoraro: 38 valves (BD 618), Petti di Carrubbare: 1 valve (BD 619), Pezzo: 6 valves (BD 620), San Procopio: 10 valves (BD 621), Saracinello: 1 valve (BD 622), Stalettì: 4 valves (BD 623), Terreti: 1 valve (BD 624), Torrente Boscaino: 6 valves (BD 625), Valle Lamato: 2 valves (BD 626); Sicily: Capo Milazzo: 11 valves (BD 627), Ficarazzi: 4 valves (BD 628), Gravitelli: 1 valve (BD 629), Messina: 1 valve (BD 630), Salice: 1 valve (BD 631), Casa Parrino: 2 valves (BD 632). Maximum width of the valves: 4.6 / 6.3 / 4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF804E620FADFBF069A19108.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight to shlightly concave. Intermediate valves broadly rectangular, width about three times length (W / L = 2.80 – 3.17), carinate in anterior profile, moderately elevated (dorsal elevation 0.33 – 0.41), anterior margin regularly convex or somewhat sinuate, side margins slightly rounded, posterior margin slightly concave at both sides of pronounced apex, lateral areas raised, Tail valve semicircular (W / L = 1.58 – 1.65), anterior margin straight or slightly convex, central mucro, small, hardly raised, antemucronal slope almost straight, postmucronal slope straight or very slightly concave directly behind mucro. Tegmentum granulose, rough. HV, LA and PMA sculptured with fine microgranules, fuse into continuous radial lines, marked with a few concentric growth lines toward outer margins, and with characteristic black dots, pigmented cusps of shell-eyes, irregularly distributed in HV and PMA, in radiating rows in LA, except for a narrow strip along posterior margin. CA and AMA sculptured with longitudinal lines of micraesthetes interrupted only by megalaesthetes irregularly distributed, with 3 – 8 longitudinal grooves strongly variable, which may reach upper edge. Articulamentum with apophyses wide, short, regularly rounded, connected at jugum by a lamina, insertion plates short, with many slits, slit formula 15 – 16 / 2 / 14 – 16, teeth irregular, slit rays well visible, eaves very porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF804E620FADFBF069A19108.taxon	discussion	Remarks. Some confusion exists about the taxonomic assessment of Callochiton doriae (Capellini, 1859) and C. septemvalvis (Montagu, 1803) (see Dell’Angelo & Smriglio 1999; Carmona Zalvide et al. 2002). Kaas (1978) proposed to separate at subspecific level the typical Atlantic (Callochiton septemvalvis septemvalvis) from the Mediterranean form (C. septemvalvis euplaeae O. G. Costa, 1829), the latter being characterized by a smaller size and the presence of longitudinal grooves on the pleural areas. Subsequently, Sigwart et al. (2013) recognize that Atlantic Callochiton septemvalvis (France) and Mediterranean Callochiton euplaeae (Croatia, Adriatic Sea) are genetically different enough to warrant their specific separation. The species were identified as Chiton euplaeae by O. G. Costa (1830, not 1829, as usually indicated, see Fasulo 2013) and Chiton doriae by Capellini (1859), respectively; however, the correct attribution to C. euplaeae is questionable, because Chiton euplaeae must be considered a nomen dubium (Dell’Angelo & Palazzi 1994; Dell’Angelo et al. 2016). We follow here Dell’Angelo et al. (2016) and attribute fossil valves of Callochiton without longitudinal grooves on the pleural areas to Callochiton septemvalvis, whilst those provided of longitudinal grooves to Callochiton doriae (Capellini, 1859) synonym of Callochiton euplaeae, nomen dubium). Already Bałuk (1971) mentioned the similarity between Chiton rariplicatus Reuss 1860 from the Paratethys and C. laevis (Montagu, 1803) (now Callochiton doriae) and later many other authors (e. g., Bałuk 1984; Macioszczyk 1988; Studencka & Studencki 1988; Ruman & Hudáčková 2015) confirm the conspecifity of Chiton rariplicatus and Callochiton doriae. All authors describe and illustrate the valves from the Paratethys with longitudinal grooves on CA and AMA, agreing with the attribution to C. doriae. Only Bałuk (1971: p. 461) reports: “ ... shallow and very distinct longitudinal furrows occur on the central area of most specimens ”, suggesting that valve without furrows (and therefore attributable to C. septemvalvis) could also be present. Considering that there are no other reports or figures of valves without grooves on CA and AMA, and with the further confirmation of all the material examined by us, we believe that only Callochiton doriae is present in Paratethys, and not C. septemvalvis. The number of longitudinal grooves is variable, between 3 and 8, mainly with short grooves along the diagonal ridge (Figs 51 F, 51 J) or with longer grooves, of which only a few reaching the anterior margin (Figs 51 C, 51 E). A tail valve from the Pliocene of Caranchi (Fig. 51 I) show the development of a new insertion plate under the already existing one, an abnormality already known but rarely reported, and especially rare in fossil valves (Dell’Angelo & Schwabe 2010). Comparisons. This species is very similar to Callochiton septemvalvis (see below), from which it differs mainly by the presence of some longitudinal grooves on CA and AMA. Other differences are discussed by Carmona Zalvide et al. (2002), in a study on Callochiton spp. living in the Iberian Peninsula (see below under Callochiton septemvalvis).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF804E620FADFBF069A19108.taxon	distribution	Distribution. Lower Miocene: Proto – Mediterranean Sea (Burdigalian): N. Italy: Sciolze (Dell’Angelo et al. 2016). Middle Miocene: Proto – Mediterranean Sea (Langhian): Po Basin, N. Italy: Albugnano (Dell’Angelo et al. 2016); Central Paratethys (Langhian – Serravallian): Austria: Steinabrunn (Šulc 1934), Czech Republich: Knínice, Rudoltice (Reuss 1860; Šulc 1934), Hungary: Bánd, Letkés (this study), Poland: Korytnica, Lychów, Monastyrz, Nawodzice Rybnica, Węglin, Weglinek (Bałuk 1965, 1971, 1984; Jakubowski & Musiał 1977; Macioszczyk 1988; Studencka & Studencki 1988), Romania: Lapugy (this study), Slovakia: Devínska Nová Ves, Kúty, Rohožník (Kováč et al. 1999; Ruman & Hudáčková 2015); Eastern Paratethys: Ukraine: Varovtsi (this study). Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place, Renauleau (Dell’Angelo et al. 2018 b); Proto – Mediterranean Sea (Tortonian): Po Basin, N Italy: Rio di Bocca d’Asino, Villa Monti (Laghi 1977; Dell’Angelo & Smriglio 1999; Dell’Angelo et al. 2016). Lower Pliocene: central Mediterranean, Italy: many localities in Liguria (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b). Pliocene: northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo & Silva 2003; Dell’Angelo et al. 2022); western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: many localities in Tuscany, Emilia-Romagna, Umbria, Sicily (Dell’Angelo et al. 2001 a; Chirli 2004; Dell’Angelo et al. 2012). Upper Pliocene to upper Pleistocene: central Mediterranean, France: Bosq d’Aubigny (Dell’Angelo et al. 2018 b), Greece: Rhodes Island (Koskeridou et al. 2009). Pleistocene: central Mediterranean, Italy: many localities in Tuscany, Emilia-Romagna, Puglia, Calabria, Sicily (Sabelli & Taviani 1979; Dell’Angelo & Giusti 1997; Dell’Angelo et al. 2001 a; this study); Greece: Kyllini (Garilli et al. 2005). Recent: Atlantic coasts of Spain near Gibraltar (Borja 1987; Carmona Zalvide & García 2000; Carmona Zalvide et al. 2002), Canary Islands (Kaas 1991) and Madeira Archipelago (Segers et al. 2009), Morocco: Cape Yubi (Kaas 1991). Mediterranean Sea: France (Leloup 1934); Italy: many localities (Leloup & Volz 1938; Dell’Angelo & Smriglio 1999); Croatia (Leloup & Volz 1938; Dell’Angelo & Zavodnik 2004); Greece and Aegean Sea Islands (Strack 1988, 1990; Zenetos & Van Aartsen 1995; Koukouras & Karachle 2005); Lebanon (Crocetta et al. 2014); Israel (Barash & Danin 1977); Algeria: Orano (Pallary 1900); Tunisia: Gulf of Gabes (Kaas 1989; Cecalupo et al. 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9D4E630FADFBEB6BBD905F.taxon	description	Fig. 52	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9D4E630FADFBEB6BBD905F.taxon	materials_examined	Type material. Holotype MNHN. F. A 67112, intermediate valve, width 4 mm (Figs 52 A – C). Paratypes: MNHN. F. A 670113 – A 67114 (2 valves, Figs 52 E – G); NHMW 2017 / 0108 / 0029 – 0030 (2 valves, Fig. 52 H); RGM. 1008411 – 1008412 (2 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): Saint-Clément-de-la-Place: type material plus 22 valves (MNHN. F. A 67115, Fig. 52 D, MNHN. F. A 67116, NHMW 2017 / 0108 / 0031, RGM. 1008413, BD 147). Maximum width of the valves: – / 6 / 2.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9D4E630FADFBEB6BBD905F.taxon	description	Description. Head valve unknown. Intermediate valves polygonal (W / L = 1.90 – 1.94), carinate in anterior profile, elevated (H / W = 0.58 – 0.64), anterior margin slightly convex in jugal area, side margins almost straight, inclined, posterior margin slightly concave either side of strongly protruding apex, lateral areas raised. Tail valve almost circular (W / L = 1.40), elevated, anterior margin convex, posterior margin almost semicircular, mucro subcentral or slightly anterior, antemucronal and postmucronal slopes almost straight, forming angle of ca. 40 °. Tegmentum granulose, rough. LA and PMA sculptured with fine granules, fusing into continuous radial lines, marked with few concentric growth lines toward outer margins. CA and AMA sculptured with longitudinal lines of micraesthetes, interrupted only by megalaesthetes irregularly distributed, with 6 – 8 conspicuous longitudinal grooves, shortening towards jugum. Articulamentum with short, wide, regularly rounded apophyses connected at jugum by lamina, insertion plates short, teeth somewhat thickened at edges of slits, slit formula – / 3 / 16, teeth irregular, slit rays visible, eaves very porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9D4E630FADFBEB6BBD905F.taxon	discussion	Remarks. The fossil record of Callochiton pouweri Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is limited to the Miocene (Tortonian) of France. The available material is not well preserved; most of the valves are small, incomplete and abraded, and head valves were not present. The presence of pigmented aesthetes (present in many species of the genus Callochiton, see Schwabe 2010; Speiser et al. 2011) has not been confirmed in C. pouweri. Comparisons. Callochiton pouweri is similar to C. calcatus Dell’Angelo & Palazzi, 1994, from which it differs by the shape of the tail valve (mucro in anterior position and an exceptionally convex postmucronal slope in C. calcatus, see above).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9D4E630FADFBEB6BBD905F.taxon	distribution	Distribution. Middle Miocene: northeastern Atlantic (Tortonian): Anjou, France: Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9C4E650FADFA2168D19164.taxon	description	Fig. 53	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9C4E650FADFA2168D19164.taxon	materials_examined	Type material. Holotype NHMUK (fide Kaas & Van Belle 1985 b). Type locality. Salcomb Bay (England). Material examined. Miocene (Tortonian): Italy: Borelli: 1 valve (BD 633, Figs 53 F – H) Rio di Bocca d’Asino: 33 valves (BD 634, MZB 32062 – 32063, PG), Montegibbio: 10 valves (BD 635, MZB 32064, Fig. 53 E). Pliocene: Portugal: Vale de Freixo: 269 valves (GeoFCUL VFX. 03.355, GeoFCUL VFX. 03.337 – 338, RGM. 1364009 – 1364011, MNHN. F. A 81986, BD 241); Spain: Estepona: 2 valves (BD 636); Italy: Poggio alla Fame: 1 valve (BD 637, Figs 53 A – C), Serre di Rapolano: 2 valves (BD 638, Fig. 53 D). Maximum width of the valves: 2 / 6.8 / 3.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9C4E650FADFA2168D19164.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight to slightly concave. Intermediate valves broadly rectangular, width about three times length (W / L = 2.86 – 3.16), carinate in anterior profile, moderately elevated (H / W = 0.35 – 0.46), anterior margin regularly convex or somewhat sinuate, side margins slightly rounded, posterior margin slightly concave at both sides of pronounced apex, lateral areas raised. Tail valve semicircular, anterior margin straight or slightly convex, central mucro small, hardly raised, antemucronal slope almost straight, postmucronal slope straight or very slightly concave directly behind mucro. Tegmentum granulose, rough. HV, LA and PMA sculptured with fine microgranules, fuse into continuous radial lines, marked with a few concentric growth lines toward outer margins, and with characteristic black dots, pigmented cusps of shell-eyes, irregularly distributed in HV and PMA, in radiating rows in LA, except for a narrow strip along posterior margin. CA and AMA sculptured with regular longitudinal lines of micraesthetes interrupted only by megalaesthetes distributed normally in a six-row recurring sequence. Articulamentum with apophyses wide, short, regularly rounded, connected at jugum by a lamina, insertion plates short, with many slits, slit formula 19 – 20 / 2 – 3 / 17 – 18, teeth irregular, slit rays well visible, eaves very porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9C4E650FADFA2168D19164.taxon	discussion	Remarks. The fossil record of Callochiton septemvalvis (Montagu, 1803) is limited to the Miocene (Tortonian) of Italy, and Pliocene of Italy, Spain and Portugal. We follow Dell’Angelo et al. (2016) by attributing fossil valves of Callochiton without longitudinal grooves on the pleural areas to Callochiton septemvalvis, whilst those provided of longitudinal grooves, to Callochiton doriae (Capellini, 1859) (synonym of Chiton euplaeae O. G. Costa, 1830, nomen dubium). We are unable to ascertain whether literature records of C. septemvalvis do all pertain to Montagu’s species, or if they encompass C. doriae. Thus, we report in the synonymy only those specimens that are safely ascribable to one or another taxon based upon the morphological details discussed above, while dubious cases are also reported in the bibliographic references, with the species preceded by a question mark. The sculpturing of the dorsal surface was highlighted by Baxter & Jones (1984) and Carmona Zalvide et al. (2002). Although past authors did not fully appreciate the characteristics differentiating this species from C. doriae, we consider C. septemvalvis as absent from the present Mediterranean chiton fauna, although its fossil record in this basin leaves the question open. Comparisons. This species is quite similar to Callochiton doriae (see above), from which it differs mainly by the absence of longitudinal grooves on CA and AA. Carmona Zalvide et al. (2002), in their study on modern Callochiton spp. of the Iberian peninsula, showed other differences regarding the maximum size of the living specimens (22 mm for C. septemvalvis vs. 18 mm for C. doriae), the structure of the aesthetes, both as regards the maximum diameter of the megalaesthetes, almost double for C. septemvalvis (10 µm vs. 5.5 µm for C. doriae), and for the average distance between the micraesthetes aligned on the longitudinal ribs (3 µm for C. septemvalvis vs. 16 µm for C. doriae); the species differs also for the slit formula (19 / 2 – 3 / 17 – 18 for C. septemvalvis vs. 15 – 16 / 2 / 14 – 16 for C. doriae), and for characters not evaluable in fossil material, i. e. the size of all three types of girdle’s (dorsal, marginal and ventral) spicules.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9C4E650FADFA2168D19164.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Rio di Bocca d’Asino, Montegibbio (Laghi 1977; Dell’Angelo et al. 2016). Pliocene: North Atlantic: Netherlands (Strack 2010); northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo & Silva 2003; Dell’Angelo et al. 2022); western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Bibbiano, Serre di Rapolano (Dell’Angelo & Forli 1995 a; this study). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece: Rhode Island (Koskeridou et al. 2009). Pleistocene: North Atlantic: Sweden and Norway (Brogger 1901; Antevs 1917, 1928). Recent: Northeastern Atlantic Ocean, from Scandinavia: N. Norway N to 67 ° N (Dons 1934; Hansson 1998) along the European coasts (McKay & Smith 1979) to Portugal and Spain (Rolan Mosquera et al. 1990; Consolado Macedo et al. 1999; Urgorri et al. 2017) and the Canary Islands (Hernández & Rolán 2011; Dell’Angelo & Smriglio 1999).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9A4E660FADFB3869FC96F8.taxon	description	Fig. 54	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9A4E660FADFB3869FC96F8.taxon	materials_examined	Type material. Holotype, unknown, not present in the Šulc collection at NHMW, intermediate valve (Šulc 1934: pl. 1, fig. 18), Fig. 54 A. Type locality. Kninice (Czech Republic). Type stage. Middle Miocene. Material examined. No actual material available, only descriptions and illustrations from the literature (Fig. 54 A – D). Maximum width of the valves: 3 / 6.5 / 6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9A4E660FADFB3869FC96F8.taxon	description	Description. Head valve semielliptical. Intermediate valves broadly rectangular, keeled, anterior margin convex, side margins slightly rounded, posterior margin almost straight with a small apex, lateral areas raised. Tail valve semielliptical, anterior margin slightly convex, mucro subcentral or slightly anterior. Tegmentum rough. HV, LA and PMA finely granulose, sculptured with zigzagging wrinkles. CA and AMA sculptured with closely and regularly spaced fine pits which radially diverge from apex. Articulamentum with apophyses narrow, widely spaced and rounded, connected at jugum by a lamina, insertion plates with many slits, slit formula 30 – 33 / 5 – 7 / 30, slit rays well visible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9A4E660FADFB3869FC96F8.taxon	discussion	Remarks. Callochiton zigzag Šulc, 1934 is a poorly known species from the Paratethys, only reported by Šulc (1934) from the Czech Republic, and by Bałuk (1971, 1984) from Korytnica (Poland). The sculpture of the valves is, however, very characteristic and different from all other Callochiton species from the European Neogene. The presence of pigmented aesthetes (present in many species of the genus Callochiton, see Schwabe 2010; Speiser et al. 2011) has not been confirmed in C. zigzag. Comparisons. Callochiton zigzag has the highest number of slits compared to the other Callochiton discussed here, and the sculpture also differs (see Tab. 8).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF9A4E660FADFB3869FC96F8.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Czech Republic: Kninice, Borač, Židlochovice (Šulc, 1934); Poland: Korytnica (Bałuk, 1971, 1984).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF994E660FADFB7C68FC9288.taxon	type_taxon	Type species. Chiton textilis Gray, 1828, by subsequent designation (Gray 1847 b: 168). For synonymy, see Kaas & Van Belle 1990.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF994E660FADFB7C68FC9288.taxon	distribution	Distribution. Ischnochiton is known from the Jurassic to the Recent, with a widespread extant circumglobal distribution, except for the northern Atlantic and Arctic Oceans (Kaas & Van Belle 1990). The fossil record extends back to the Jurassic of Germany (Fiedel & Keupp 1988), the Eocene in Europe (U. K. and Ukraine: Bielokrys 1999; Cherns & Schwabe 2019), the upper Eocene or lower Oligocene in Washington, U. S. A. (Dell’Angelo et al. 2011 a), the Oligocene of France (Dell’Angelo et al. 2020 b) and New Zealand (Lee et al. 2014; Wu & Lee 2024), the Miocene of Africa (Tanzania: Davis 1954), Indonesia (Sumatra: Van der Vlerk 1931), the Miocene-Pleistocene of Europe (Dell’Angelo et al. 2004, 2015 a; Garilli et al. 2005; Studencka & Dulai 2010), Australia and New Zealand (Cotton 1964; Beu & Maxwell 1990), Pliocene-Pleistocene of U. S. A. (Berry 1940; Campbell 1993), the Pleistocene of Japan (Itoigawa et al. 1976), Uruguay (Rojas & Urteaga 2011) and Red Sea (Dell’Angelo et al. 2020 a), the Holocene of Japan (Kuroda et al. 1980).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF994E660FADFB7C68FC9288.taxon	discussion	Remarks. The genus Ischnochiton shows a high number of species treated in this work, 17 in all, reported by source area (Mediterranean, NE Atlantic and Paratethys), while for a better identification and differentiation, the examined species are reported, in the tables that summarize the main diagnostic characters, in two groups which highlight the different tegmentum sculpture:	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF934E6D0FADFF286FFC94B4.taxon	description	Fig. 55	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF934E6D0FADFF286FFC94B4.taxon	materials_examined	Type material. Holotype: MPUM 18956 (1 intermediate valve). Paratypes: MPUM 18957 (1 head and 1 intermediate valves); MPUM 18958 (7 intermediate valves). Type locality. Tagliata, Modena (Emilia-Romagna, Italy). Type stage. Lower Pliocene. Material examined. Pliocene: Italy: Tuscany: Luciana: 2 valves (BD 441), Pietrafitta: 1 valve (BD 442, Figs 55 D – E); Emilia-Romagna: Cava di Campore: 5 valves (BD 443, Figs 55 F – H), Torrente Stirone: 2 valves (BD 444); Sicily: Altavilla: 9 valves (AG, AR, BD 445, Figs 55 A – C). Maximum width of the valves: 4.7 mm (intermediate valves).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF934E6D0FADFF286FFC94B4.taxon	description	Description. Head and tail valve unknown. Intermediate valve broadly rectangular (W / L = 2.66 – 2.75), carinate in anterior profile, highly elevated (H / W = 0.48 – 0.52), anterior border consisting of three segments, the two lateral ones straight and the one between the apophyses almost straight to slightly convex, side margins almost straight to slightly convex, posterior margin nearly straight, with clearly prominent apex, lateral areas moderately elevated. LA weakly sculptured with a fine granularity giving a smooth appearance, with many concentric grooves. CA with reticulate sculpture consisting of obliquely intersecting vermicular ribs, with granules tending to separate in JA, and an almost smooth area, with some megalaesthetes arranged in parallel rows, present towards side margins. Many aesthetes irregularly distributed on tegmentum’s surface, some arranged in parallel rows present in smooth area. Articulamentum with apophyses wide, triangular, 2 slits each side.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF934E6D0FADFF286FFC94B4.taxon	discussion	Remarks. The fossil record of Ischnochiton anserinus Laghi, 1977 is limited to the Italian Pliocene, and is extended here to few other localities (Cava di Campore, Torrente Stirone). This rare species is characterized by the distinctly carinated (like the breastbone of a goose, hence the name of the species) intermediate valves, and the tegmentum sculptured with a fine granularity formed by a trellis-work of diagonal furrows, well evidenced in JA, giving a smooth appearance on LA and PA towards the side margins. Laghi established his new species on the basis of nine poor preserved and incomplete intermediate valves, and only a small fragment of a head valve, with the same sculpture of a fine granularity (practically not visible on the Laghi’s figure) giving a smooth appearance. All subsequent records of I. anserinus refer to intermediate valves. Comparisons. This species is somewhat related to I. martinelli Dell’Angelo, Landau & Marquet, 2004; the differences are discussed under the latter species.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF934E6D0FADFF286FFC94B4.taxon	distribution	Distribution. Pliocene: central Mediterranean, Italy: Tuscany: Luciana, Pietrafitta; Emilia-Romagna: Cava di Campore, Torrente Stirone and Sicily: Altavilla Milicia (Laghi 1977; Dell’Angelo et al. 2001 a, 2012; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF924E6E0FADFE486B5F9724.taxon	description	Fig. 56	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF924E6E0FADFE486B5F9724.taxon	materials_examined	Type material. Holotype MZB 50513 (tail valve, width 5.3 mm, Figs 56 E – H). Paratypes: MZB 50514 (head valve, Figs 56 A – B), MZB 50515 (intermediate valve, Fig. 56 C). Type locality. Archi S. Francesco, Reggio Calabria (Calabria, Italy). Type stage. Upper Pleistocene. Material examined. Lower Pleistocene: Italy: Archi, Fornace Aloi (lower Pleistocene): 1 valve (BD 446). Upper Pleistocene: Italy: Archi S. Francesco: type material plus 19 valves (BD 447, Fig. 56 D). Maximum width of the valves: 6.8 / 5.5 / 5.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF924E6E0FADFE486B5F9724.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight. Intermediate valves broadly rectangular, anterior margin slightly convex, side margins rounded, posterior margin straight, apex inconspicuous, lateral areas not raised, separated from central area by different sculpture. Tail valve semicircular (W / L = 1.71), anterior margin slightly convex, mucro subcentral, swollen but not prominent, antemucronal slope slightly convex, postmucronal slope slightly concave just behind mucro. HV, LA and PMA sculptured with radial nodulose riblets (HV 34 – 42, 34 in paratype; LA 5; PA 38 in holotype), some of which tending to split near outer margins, roundish to squarish nodules become larger towards periphery. CA and AMA sculptured with nodulose longitudinal riblets (CA ca. 20 – 30 on half valve, some more irregular near side margin; AMA 42 in holotype), with granules come nearer to each other, not distinctly separated. Articulamentum with teeth irregular, slit formula: 12 / 1 / 9, slit rays well visible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF924E6E0FADFE486B5F9724.taxon	discussion	Remarks. The fossil record of Ischnochiton crovatoi Dell’Angelo, Sosso & Bonfitto, 2019 is limited to the Pleistocene of S. Italy. The material examined is in some case not well preserved or complete (especially the intermediate valves, none of them complete), and show a certain variability regarding the number of radial nodulose riblets on HV, ranging between 34 and 42, counting only the riblets without splitting (ca. 37 – 47 counted on the anterior margin). Comparisons. The characteristic ornamentation of Ischnochiton crovatoi, formed by radial nodulose riblets on HV, LA and PMA, distinguishes it from the common Mediterranean species I. rissoi (Payraudeau, 1826); in this latter species the sculpture consisting of concentric wavy lirae often crossed by fine radiating grooves, that gives to the tegmentum a granular outlook.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF924E6E0FADFE486B5F9724.taxon	distribution	Distribution. Pleistocene: central Mediterranean, Italy: Archi S. Francesco, Archi Fornace Aloi (Dell’Angelo et al. 2019 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	description	Fig. 57	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	materials_examined	Type material. Holotype: MZB 49983, an intermediate valve from Bussana (Figs 57 B – D). Paratypes: MZB 49981 – 49982 (tail and head valves from Caranchi and Zinola, Fig. 57 A); MSNG 56536 (intermediate valve from Rio Torsero); MGPT PU 109800 – 109801 (intermediate and tail valves from Rio Torsero and Caranchi), and private collections: BD (3 valves, 2 from Bussana and 1 from Zinola), MS (1 tail valve), MP (1 tail valve from Rio S. Antonino). Type locality. Bussana (Liguria, Italy). Type stage. Lower Pliocene (Zanclean). Material examined. Upper Miocene: Italy: Montegibbio: 2 valves (BD 448, MZB 32051), Rio di Bocca d’Asino: 3 valves (BD 449, MZB 32028 – 32029). Lower Pliocene: Italy: Liguria: type material plus Borzoli: 9 valves (BD 450, Figs 57 G – H), Bussana: 20 valves (BD 451); Caranchi: 5 valves (BD 452, MP), Garlenda: 20 valves (MP), Genova Sestri: 4 valves (BD 453), Rio Sant’ Antonino: 40 valves (MP, Fig. 57 E), Rio Torsero: 2 valves (BD 454, Fig. 57 F), Zinola: 4 valves (BD 455). Maximum width of the valves: 3.2 / 3.8 / 3.6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight. Intermediate valve broadly rectangular (W / L = 2.34 – 2.50), anterior margin straight, moderately elevated (H / W = 0.33), rounded to semicarinate in anterior profile, side margins slightly rounded, posterior margin practically straight, apex not very visible, lateral areas scarcely differentiated. Tail valve semicircular (W / L = 1.68 – 1.98), anterior margin almost straight, mucro central, not prominent, antemucronal slope practically straight, postmucronal slope slightly concave just behind mucro. Tegmentum uniformly sculptured with very irregular granules, arranged in segments of various size and shape, slightly overlapping each other, forming rugosities. HV, LA and PMA sculptured with some rugosities acquiring vaguely concentrical zig-zag pattern, producing undulating macrosculpture. CA and AMA sculptured with rugosities sometimes giving impression of very irregular longitudinal chains of granules, or sinuose and intersecting each other, as a network of irregular, elongate pits. Each granule with many aesthetes of same size irregularly distributed. Articulamentum with large apophyses, teeth of irregular width, deeply incised, slit formula 12 / 1 / 10 – 12.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	discussion	Remarks. The fossil record of Ischnochiton ligusticus Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013 is limited to the Miocene (Tortonian) of N. Italy (Po Basin) and the Pliocene (Zanclean) of N. Italy (Liguria). The studied material is represented by valves sometimes incomplete and poorly preserved, especially the characters of the articulamentum are scarcely visible. Comparisons. Ischnochiton ligusticus is superficially similar to Ischnochiton ulivii Dell’Angelo & Forli, 1996, from which it differs mainly by the very irregular sculpture, while I. ulivii is by contrast characterized by small granules quincuncially arranged on HV, LA and PMA, and by a decidedly pitted sculpture on CA and AMA.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	distribution	Distribution: Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, North Italy: Montegibbio, Rio di Bocca d’Asino (Dell’Angelo et al. 2015 a). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Caranchi, Garlenda, Genova Sestri, Rio S. Antonino, Rio Torsero, Zinola (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b). “ Ischnochiton ” luquei Dell’Angelo, Sosso & Taviani, 2024 Fig. 58	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	materials_examined	Type material. Holotype: MZUB 60343, intermediate valve, width 2.5 mm (Figs 58 A – D). Paratype 1: (MZUB 60344), intermediate valve, width 2.8 mm; Paratype 2: MZUB 60345, intermediate valve, width 2.5 mm; Paratype 3: MZUB 60346, intermediate valve, width 2.8 mm; Paratype 4 (MNHN-IM- 2022 - 2405), intermediate valve, width 3.1 mm; Paratype 5 SMF 376793, intermediate valve, width 2.8 mm; Paratype 6 (MZUB 60347), intermediate valve, width 3 mm. Type locality. Capraia Island-Capo Corso, sediments – 350 / 500 m by fishermen. Type stage. Pleistocene, presumably last glacial. Material examined. Pleistocene, presumably last glacial: Italy: Tuscany: Capraia Island-Capo Corso - 350 / 500 m: type material plus 9 intermediate valves (BD 251). Maximum width of the valves: 3.4 mm (intermediate valves).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	description	Description. Head and tail valve unknown. Intermediate valve broadly rectangular (W / L = 1.72 – 1.96), semicarinate in anterior profile, elevated (H / W = 0.45 – 0.50), anterior margin convex, side margins almost straight or little rounded, posterior margin almost straight at both sides of protruding apex, lateral areas raised. Tegmentum smooth. LA smooth with some weak concentric growth lines. CA smooth, with some weak concentric growth lines, continuing from LA. Articulamentum with large, triangular apophyses, insertion plate short with 1 slit, slit rays well visible, a second slit ray is almost always visible near posterior margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	discussion	Remarks. “ Ischnochiton ” luquei Dell’Angelo, Sosso & Taviani, 2024 is only known from the Tuscan Archipelago, Mediterranean Sea, offshore Capraia Island-Capo Corso at - 350 / 500 m. Only intermediate valves are known to date. Also, the generic attribution is difficult, many extant species with a smooth or microgranulose tegmentum are attributed to the genera Ischnochiton Gray, 1847, Stenosemus von Middendorff, 1847 or Lepidochitona Gray, 1821, mainly based upon the soft parts. We attribute provisionally the material studied to Ischnochiton. Comparisons. “ Ischnochiton “ luquei is superficially similar to Boreochiton ruber (Linnaeus, 1767), a species with an arctic-circumboreal distribution (Kaas & Van Belle 1985; Sirenko & Dell’Angelo 2023), with whom it shares the smooth tegmentum, and from which it differs by the different shape of the intermediate valves (much more elongate in B. ruber), and the weaker concentric growth lines, more prominent in B. ruber.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF914E900FADFDF868379126.taxon	distribution	Distribution. Pleistocene: central Mediterranean, Italy: Capraia Island-Capo Corso, sediments 350 / 500 m (Dell’Angelo, Sosso & Taviani 2024).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6F4E910FADFBFE685392D6.taxon	description	Fig. 59	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6F4E910FADFBFE685392D6.taxon	materials_examined	Type material. Holotype: MZB 25051 (intermediate valve, Figs 59 C – D). Paratypes: MZB 25052 (3 valves); MME (3 valves, Figs 59 A – B); IRSN IST 6450 (3 valves); BD 4567 (3 valves); RM (3 valves). Type locality. Estepona, Velerín Carretera (Málaga, Spain). Type stage. Early Piacenzian (Pliocene). Material examined. Lower Pliocene: Italy: Liguria: Caranchi: 2 valves (BD 456, MZB 45716), Rio Sant’Antonino: 1 valve (MP). Pliocene: Spain: Estepona: type material plus 26 valves (BD 457). Italy: Emilia-Romagna: Cava di Campore: 8 valves (BD 458), Lugagnano Val d’Arda: 1 valve (BD 459); Tuscany: Orciano Pisano: 1 valve (BD 460); Sicily: Altavilla: 24 valves (AG, AR, BD 461, Figs 59 E – F). Pleistocene: Italy: Calabria: Archi S. Francesco: 35 valves (BD 462, Figs 59 G – L), Bovetto: 1 valve (BD 463), Terreti, Ponte Cellantoni: 4 valves (BD 464); Sicily: Salice: 3 valves (BD 465). Maximum width of the valves: 4.8 / 6.5 / 4.1 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6F4E910FADFBFE685392D6.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, notched in middle, front slope straight to slightly concave. Intermediate valves broadly rectangular (W / L = 2.57 – 3.01), carinate in anterior profile, moderately to highly elevated (H / W = 0.36 – 0.54), anterior margin consisting of three segments, two lateral segments concave, and the one between the apophyses convex, side margins slightly convex, posterior margin nearly straight, apex little evident, lateral areas moderately elevated. Tail valve semicircular (W / L = 2.10), with anterior margin similar to that of intermediate valves, mucro subcentral, antemucronal slope slightly convex, postmucronal slope slightly concave behind mucro HV, LA and PMA sculptured with radial nodulose riblets (HV 26 – 30, LA 4 – 5), becoming indistinct toward apex, with squarish nodules formed by many concentric grooves crossing riblets. CA and AMA sculptured with a reticulate sculpture consisting of obliquely intersecting vermicular ribs, with granules tending to separate in JA, and tending to form about ten real longitudinal striae in pleural area, often somewhat eroded, and an almost smooth area, with some megalaesthetes arranged in parallel rows, present towards side margins. Tegmentum fully covered by aesthetes, both on nodular elevations (without a clearly recognizable structure of megalaesthetes and micraesthetes) and between them. Articulamentum with apophyses wide, triangular in intermediate valves, trapezoidal in tail valve, separated by narrow jugal sinus, slit formula 9 – 11 / 1 – 2 / 11, slits inequidistant.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6F4E910FADFBFE685392D6.taxon	discussion	Remarks. The fossil record of Ischnochiton martinelli Dell’Angelo, Landau & Marquet, 2004, so far limited to the Pliocene of Estepona (Spain) and Italy (Liguria and Sicily) is extended here to other Pliocene and Pleistocene localities from Italy. The valves from the Italian Pleistocene well agree with the same material from the Pliocene localities. The number of slits in the intermediate valves is more difficult to evaluate, also because the characteristics of the articulamentum are not always clearly visible. Some valves show a second slit ray near the posterior margin, but without evidence of a real slit, which is instead clearly noticeable in the valve illustrated (Fig. 54 I) from Archi S. Francesco. Comparisons. Ischnochiton martinelli is similar to I. anserinus Laghi, 1977, only known from intermediate valves, with which it shares the carinate anterior profile of intermediate valves, and from which it differs mainly by the sculpture of LA, with a fine granularity, giving a smooth appearance in I. anserinus, vs. radial nodulose riblets with squarish nodules in I. martinelli.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6F4E910FADFBFE685392D6.taxon	distribution	Distribution. Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Altavilla, Caranchi, Rio S. Antonino (Dell’Angelo et al. 2012, 2013), Cava di Campore, Lugagnano Val d’Arda, Orciano Pisano (this study). Pleistocene: central Mediterranean, S. Italy: Archi S. Francesco, Bovetto, Salice, Terreti (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6D4E940FADFF056BED92E0.taxon	description	Fig. 60	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6D4E940FADFF056BED92E0.taxon	materials_examined	Type material. Syntype MNHN 6109. Type locality. Bonifacio (Corse, France). Material examined. Middle Miocene: Central Paratethys: Austria: Grund: 1 valve (NHMW 1868 / 0001 / 0281), Pötzleinsdorf: 18 valves (NHMW 1859 / 0027 / 0062, Figs 60 S – T, 1859 / 0038 / 0190, 2010 / 0256 / 0007); Romania: Kostej: 2 valves (BD 466, NHMW 2010 / 0256 / 0003), Lapugy: 3 valves (BD 467); Hungary: Bánd: 3 valves (BD 468), Letkés: 4 valves (BD 469); Eastern Paratethys: Ukraine: Horodok: 3 valves (BD 470), Varovtsi: 133 valves (BD 471, Figs 60 M – R). Upper Miocene (Tortonian): France: Saint-Clément-de-la-Place: 7 valves (MNHN. F. A 67094, RGM. 1008373, Figs 60 K – L, RGM. 1008374, BD 140). Italy: Borelli: 1 valve (MGPT PU 135040), Montegibbio: 4 valves (BD 472, MZB 32019), Rio di Bocca d’Asino: 12 valves (BD 473, PG, MZB 32018, MZB 32020, Figs 60 E – F), Villa Monti: 2 valves (BD 474, MZB 32049). Lower Pliocene: Italy: Liguria: Borzoli: 7 valves (BD 475), Bussana: 4 valves (BD 476, PG, MZB 45707, MZB 45709), Garlenda: 1 valve (MP), Rio Sant’Antonino: 26 valves (BD 477, MP), Rio Torsero: 6 valves (BD 478, PG, SR, MZB 45708). Pliocene: Spain: Estepona: 13 valves (BD 479). Italy: Emilia-Romagna: Gagliardella “ Tagliata ”: 2 valves (BD 480), Marzeno: 1 valve (BD 481), Rio Merli: 1 valve (BD 482); Tuscany: Bibbiano: 1 valve (BD 483), Castell’Anselmo: 1 valve (BD 484), Castelnuovo Berardenga: 1 valve (BD 485), Cetona: 1 valve (BD 486), Colle Val d’Elsa: 3 valves (BD 487), Montenero: 1 valve (BD 488), Orciano Pisano: 3 valves (BD 489), Pietrafitta: 46 valves (BD 490), Pietrafitta Melograni: 17 valves (BD 491), Serre di Rapolano: 29 valves (BD 492, Figs 60 B – D); Latium: Magliano Sabina: 7 valves (BD 493); Sicily: Altavilla: 3 valves (AG, AR, BD 494), Sciacca: 2 valves (BD 495), Trappeto: 6 valves (BD 496). Pleistocene: Italy: Emilia-Romagna: Torrente Stirone: 4 valves (BD 497); Tuscany: Cisternino: 2 valves (BD 498), Fauglia: 11 valves (BD 499), Riparbella: 3 valves (BD 500); Puglia: Gallipoli: 4 valves (BD 501), Marina di Novaglie: 1 valve (BD 502); Calabria: Archi S. Francesco: 50 valves (BD 503, Figs 60 A, 60 G – J), Carrabbati: 1 valve (BD 504), Castellace: 1 valve (BD 505), Le Castella: 2 valves (BD 506), Musalà: 5 valves (BD 507), Pecoraro: 8 valves (BD 508), Pezzo: 7 valves (BD 509), Saracinello: 4 valves (BD 510), Stalettì: 7 valves (BD 511), Terreti: 1 valve (BD 512), Valle Lamato: 1 valve (BD 513); Sicily: Capo Milazzo: 4 valves (BD 514), Grammichele: 1 valve (BD 515), Melilli: 1 valve (BD 516), Menfi: 2 valves (BD 517). Greece: Kyllini: 5 valves (BD 518, DGUP). Maximum width of the valves: 7.5 / 10 / 6.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6D4E940FADFF056BED92E0.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.25 – 3.12), moderately to highly elevated (H / W = 0.38 – 0.50), regularly rounded in anterior profile, anterior margin almost straight on jugal side, side margins slightly rounded, posterior margin straight, without trace of apex, lateral areas moderately elevated. Tail valve semicircular (W / L = 1.39 – 1.74), mucro centrally swollen but not prominent, antemucronal slope almost straight to slightly convex, postmucronal slope slightly concave just under mucro. Tegmental sculpture varies from obsolete to quite strongly marked. HV, LA and PA sculptured with concentric vermicular ribs, often intersected by fine radial furrows, which give an aspect of granulose radial ribs; CA and AA sculptured with quite regular longitudinal ribs, more spaced near lateral margins, thinner and closer together in jugal area. Tegmentum fully covered by aesthetes, both on nodular elevations (without a clearly recognizable structure of megalaesthetes and micraesthetes) and between them. Articulamentum with apophyses rounded, trapezoidal in tail valve, jugal sinus wide, about a third of valve’s width, insertion lamina divided into rather large, irregular teeth, slit formula: 10 – 13 / 1 / 9 – 12.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6D4E940FADFF056BED92E0.taxon	discussion	Remarks. Ischnochiton rissoi (Payraudeau, 1826) is rather common in the Mediterranean Sea, very variable in size, sculpture and colour, and this morphological variability has produced taxonomic discrepancies, with the description of numerous species or varieties, now considered as synonyms, e. g., Chiton mediterraneus Reeve, 1847, Chiton meneghinii Capellini, 1859, Lepidopleurus meneghinii var. dautzenbergi Ancey, 1898 (Kaas & Van Belle 1990; Dell’Angelo & Smriglio 1999; Carmona Zalvide & García 1999). Based upon only shell’s morphological features, the variability observed in I. rissoi mainly concerns the tegmentum sculpture in HV, LA and PA, with more or less regular radial ribs sometimes interrupted by concentric ribs, or with the vermicular ribs in LA tending to become longitudinal and parallel ribs in CA (Nierstrasz 1906: fig. 13; Carmona Zalvide & García 1999, figs 2 – 5). In the paleontological record, valves attributable to Ischnochiton rissoi are generally uncommon, with greater frequency of valves with a radial ribbing sculpture stronger than concentric ribbing on HV, LA and PA (Figs 60 D – F, 60 K, 60 M – N, 60 Q, 60 S); also the other types of sculpture are present, namely shells with radial ribs that intersect the concentric ribs showing a more granular structure (Figs 60 C, 60 G) and concentric vermicular ribs (Figs 60 A – B, 60 J) Already Šulc (1934) mentioned the similarity between I. rudolticensis Šulc, 1934 and 1. rissoi and later Laghi (1977) placed Šulc’s species into the synonymy of I. rissoi. In some earlier papers Malatesta (1962) and Bałuk (1971) supposed that I. rudolticensis is the Miocene ancestor of the Recent I. rissoi. The conspecificity of I. rudolticensis and 1. rissoi was confirmed by several authors (Bałuk 1984; Dulai 2005, Dell’Angelo et al. 2007 a; Studencka & Dulai 2010) and we agree and illustrate some material from the Paratethys (Figs 60 M – T). The valves examined show a high degree of variability in shape and sculpture, both in intermediate valves (W / L = 2.25 – 3.12, while the profile is always rounded, H / W = 0.38 – 0.50) and in tail ones (W / L = 1.39 – 1.74). Also the number of longitudinal ribs in CA and AMA is variable, the ribs are quite regular and parallel, sometimes slightly slanting towards the side margins, splitting more frequently near the anterior margin. Comparisons. The species is similar to Ischnochiton lesporti sp. nov. and I. sigwartae sp. nov.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6D4E940FADFF056BED92E0.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian – Serravallian): Austria: Gainfarn, Grund, Pötzleinsdorf (Kroh 2002, 2003; this study), Czech Republic: Kninice, Rudoltice (Šulc 1934), Hungary: Bánd, Devecser, Letkés, Várpalota (Dulai 2005, 2025 a, 2025 b; Dulai & Katona 2024; this study), Poland: Korytnica (Bałuk 1971, 1984), Romania: Kostej, Lapugiu (Dell’Angelo et al. 2007 a; this study); Eastern Paratethys: Ukraine: Horodok, Varovtsi (Studencka & Dulai 2010; this study). Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b); Proto – Mediterranean Sea (Tortonian and Messinian): Po Basin, N Italy: Borelli, Montegibbio, Rio di Bocca d’Asino, Villa Monti (Dell’Angelo et al. 1999, 2015 a). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Garlenda, Rio S. Antonino, Rio Torsero (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b). Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: many localities in Tuscany, Emilia-Romagna, Latium and Sicily (Laghi 1977; Dell’Angelo et al. 2001 a, 2012; Chirli 2004; this study). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece: Rhodes (Koskeridou et al. 2009). Pleistocene: central Mediterranean, Italy: many localities in Tuscany, Emilia-Romagna, Puglia, Calabria, Sicily (Sabelli & Taviani 1979; Dell’Angelo et al. 2001 a; this study). Middle to upper Pleistocene: central Mediterranean, Greece (Garilli et al. 2005), Italy (this study). Recent: Atlantic coasts, from Spain and Portugal (Consolado Macedo et al. 1999; Urgorri et al. 2017) south to Morocco (Pallary 1920), Canary Islands (Kaas 1991; Rolan 2011) and Berlengas Archipelago (Pisani Burnay 1986). Mediterranean Sea: France (B. D. D. 1882); Italy (Dell’Angelo & Smriglio 1999), Croatia (Dell’Angelo & Zavodnik 2004), Rhodes, Cyprus, Aegean Sea (Zenetos & Van Aartsen 1995; Koukouras & Karachle 2005), Turkey (Ozturk et al. 2014), Israel (Barash & Danin 1977; Barash 1982), Tunisia: Gulf of Gabes (Cecalupo et al. 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	description	Fig. 61	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	materials_examined	Type material. Holotype: MSNG 62639, tail valve, width 3.6 mm (Figs 61 I – K). Paratype 1: MSNG 62640, head valve, width 4.6 mm (Figs 61 A – C). Paratype 2: MSNG 62641, intermediate valve, width 4.2 mm (Figs 61 D – F). Paratype 3: MNHN. F. A 98470, intermediate valve, width 3 mm (Figs 61 G – H). Paratype 4: MNHN. F. A 98471, tail valve, width 3 mm (Fig. 61 L). Paratype 5: SMF 380819, intermediate valve, width 3 mm). Type locality. Archi S. Francesco, Reggio Calabria (Calabria, Italy). Type stage. Upper Pleistocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	etymology	Etymology. The specific name honors Julia Sigwart (Senckenberg Research Institute and Museum Frankfurt and Queen’s University Belfast), for her prominent contributions to the study of Recent and fossil chitons.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	materials_examined	Material examined. Upper Pleistocene: Italy: Archi S. Francesco: type material plus 17 valves (BD 519). Maximum width of the valves: 4.6 / 5.8 / 4.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, moderately elevated, rounded, without trace of apex, Tail valve semi-circular, mucro centrally swollen but not prominent. Tegmentum sculptured with concentric, large and irregular ribs in HV, LA, PMA, continuing longitudinally in CA and AMA, someone joined together, others splitting, tegmentum fully covered by aesthetes, both on nodular elevations and between them. Articulamentum with apophyses rounded, slit formula: 12 / 2 / 9 – 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular, moderately elevated (H / W = 0.28 – 0.37), rounded in anterior profile, anterior margin almost straight, side margins slightly rounded, posterior margin straight, without trace of apex, lateral areas not very elevated. Tail valve semicircular, mucro centrally swollen but not prominent, antemucronal slope slightly convex, postmucronal slope slightly concave. HV, LA and PMA sculptured with concentric, large and irregular ribs, very close to each other; ribs continue longitudinally on CA and AMA, more irregular, someone joined together, others splitting in various areas, mainly near anterior margin and towards LA. Tegmentum fully covered by aesthetes, both on nodular elevations (without a clearly recognizable structure of megalaesthetes and micraesthetes) and between them. Articulamentum with apophyses rounded, trapezoidal in tail valve, jugal sinus wide, insertion lamina divided into irregular teeth, slit formula: 12 / 2 / 9 – 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	discussion	Remarks. The fossil record of Ischnochiton sigwartae sp. nov. is limited to the upper Pleistocene of Archi S. Francesco (Calabria, Italy). The material is quite well preserved, although all the intermediate valves are somewhat incomplete. Comparisons. Ischnochiton sigwartae sp. nov. is similar to I. lesporti sp. nov., from which it shares the sculpture of the valves, but differs by the more flattened intermediate valves (H / W = 0.28 – 0.37 vs. 0.40 – 0.51 in I. lesporti sp. nov.), the slit formula (12 / 2 / 9 – 10 vs. 12 – 13 / 2 / 12 – 14 in I. lesporti sp. nov.), and the different geographic / stratigraphic distribution (upper Pleistocene of S. Italy vs. lower / Middle Miocene of the Aquitaine Basin for I. lesporti sp. nov.).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF6A4E960FADFF056E839794.taxon	distribution	Distribution. Upper Pleistocene: central Mediterranean, Italy: Archi S. Francesco (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF694E970FADFD686E609348.taxon	description	Fig. 62	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF694E970FADFD686E609348.taxon	materials_examined	Type material. Holotype: MZB 11654, 1 intermediate valve (Dell’Angelo et al. 2001 a). Paratypes: MZB 12698 – 12699 (1 head and 1 tail valves); DSTUT B-P. I. 07.1 (3 valves: 2 from Pietrafitta “ Sbarra ”, 1 tail from Pietrafitta Melograni); private collections: BD F 55 D / 2 (3 valves from Pietrafitta Melograni), MF (3 valves from Pietrafitta Melograni), EU (3 valves from Pietrafitta Melograni), VB F 1002 a (3 valves from Pietrafitta Melograni). Type locality. Pietrafitta “ Sbarra ” (Tuscany, Italy). Type stage. Pliocene. Material examined. Pliocene: Spain: Estepona: 1 valve (BD 520, Figs 62 K – L). Italy: Tuscany: Aiano: 1 valve (BD 521), Bibbiano: 8 valves (BD 522), Colle Val d’Elsa: 2 valves (BD 523), Orciano Pisano: 1 valve (BD 524), Pietrafitta: type material plus 54 valves (BD 525, Figs 62 A – G), Poggio alla Fame: 26 valves (BD 526). Pleistocene: Italy: Tuscany: Fauglia: 8 valves (BD 527, Figs 62 H – J), Riparbella: 2 valves (BD 528). Maximum width of the valves: 2.7 / 3.3 / 2.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF694E970FADFD686E609348.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight. Intermediate valves broadly rectangular, moderately elevated (H / W = 0.32 – 0.45), rounded in anterior profile, anterior margin straight, side margins slightly rounded, posterior margin almost straight, apex not evident or hardly pronounced in some valves, lateral area not evident, practically indistinguishable from central area. Tail valve semicircular, anterior margin straight, mucro hardly pronounced and visible, subcentral or slightly anterior, antemucronal slope almost straight or slightly convex, postmucronal slope almost straight or slightly concave. Tegmentum evenly granulose. HV, LA and PMA sculptured with minute tubercles irregularly arranged (more or less quincuncially), sometimes arranged in concentric arcs in LA, especially towards side margin, giving the impression of more or less continuous cords. CA and AMA sculptured with small but neat irregularly arranged depressions, giving a pitted appearance. Articulamentum with rather pronounced apophyses, insertion lamina divided into short and well-marked teeth, of fairly regular length, slit formula 10 – 11 / 1 – 2 / 10 – 12.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF694E970FADFD686E609348.taxon	discussion	Remarks. The fossil record of Ischnochiton ulivii Dell’Angelo & Forli, 1996 is limited to the Pliocene of Spain (Estepona) and Italy (random records in Tuscany) and the Pleistocene of Italy (Riparbella). Ischnochiton ulivii is characterized by the small dimensions, the minute tubercles more or less quincuncially arranged on the entire valve’s surface, and a decidedly pitted sculpture in CA and AMA. The tegmentum sculpture of Ischnochiton ulivii is very characteristic, and at first look resembles the four Recent species known from the Cape Verde Archipelago characterized by an evenly granulose tegmentum [I. cessaci (de Rochebrune, 1881) also from Mauritania, Sénégal, Angola, and Canary Islands, I. goreensis Thiele, 1909 also from Sénégal, I. paessleri Thiele, 1909, and I. nicklesi Kaas & Van Belle, 1990 also from Dakar], from which it differs by its smaller dimensions, the different slit formula, the shape of the tail valve, and mainly a much coarser sculpture. Comparisons. The present species is closest to Ischnochiton ligusticus Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013 (see above).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF694E970FADFD686E609348.taxon	distribution	Distribution. Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Tuscany: Aiano, Bibbiano, Colle Val d’Elsa, Orciano Pisano, Pietrafitta, Poggio alla Fame (Dell’Angelo & Forli 1996; Dell’Angelo et al. 2001 a; Chirli 2004; this study). Pleistocene: central Mediterranean, Italy: Fauglia, Riparbella (Dell’Angelo et al. 2001 a; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF674E980FADFF056BBC92D6.taxon	description	Fig. 63	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF674E980FADFF056BBC92D6.taxon	materials_examined	Type material. Holotype, ZSM Mol 20071427, intermediate valve, width 9.2 mm (Figs 63 F – H). Type locality. Abbesse (Aquitaine, France), see rectification for the locality in Dell’Angelo et al. (2018 a: 26, not Abbesse, Bourgogne). Type stage. Upper Oligocene (Chattian). Material examined. Lower Miocene: France (Burdigalian): Gamachot: MHNBx 2017.9.2, intermediate valve, width 7.8 mm (Figs 63 A – E).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF674E980FADFF056BBC92D6.taxon	description	Description. Head and tail valves unknown. Intermediate valve broadly rectangular, large, broad and short (W / L = 2.74 – 3.60), gently rounded in anterior profile, moderately elevated (H / W = 0.30 – 0.34), anterior margin straight across jugal area, rounding convexly outwards, side margins rounded, posterior margin nearly straight, narrow triangular, slightly elevated rounded jugal fold, lateral areas narrow, delineated by low diagonal beaded ridge and change of ornament. Tegmental sculpture consisting of coarse beaded ribs strong outside worn JA, longitudinal on CA, radial on LA. Apophyses short, barely projecting beyond the anterior margin, rounded, well separated across broad jugal sinus, ventral surface smooth, short wide ribbon-like apical area across posterior margin, wider and anteriorly triangular in apical region, no insertion plates visible but indications of slit rays.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF674E980FADFF056BBC92D6.taxon	discussion	Remarks. Ischnochiton abbessi (Cherns & Schwabe, 2019) is known only for two intermediate valves, the first originally described from the Oligocene of Abbesse (as Chaetopleura) and the second from the Burdigalian of Gamachot by Dell’Angelo et al. (2018 a), which assigned the valves to the genus Ischnochiton, on the basis of strong similarities to species included in this genus. The characters of the valve from the Burdigalian of Gamachot agree with those from the late Oligocene of Abbesse, with only some slight differences in the shape (W / L = 2.74 vs. 3.60, and H / W = 0.34 vs. 0.30), but the studied material is really scarce and does not allow to define the intraspecific variability of this species. Comparisons. See Tabs 9, 10 for a comparison with the Ischnochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF674E980FADFF056BBC92D6.taxon	distribution	Distribution. Upper Oligocene: northeastern Atlantic (Chattian): Aquitaine Basin, France: Abbesse (Cherns & Schwabe 2019). Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Gamachot (Dell’Angelo et al. 2018 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	description	Fig. 64	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	materials_examined	Type material. Holotype MSNG 62642, intermediate valve, width 4.6 mm (Figs 64 D – F). Paratype 1, MSNG 62643, tail valve, width 3.5 mm (Figs 64 I – L). Paratype 2, MHNBx 2024.2.1, head valve, width 3.6 mm (Figs 64 A – C). Type locality. Gaas (France). Type stage. Lower Oligocene (Rupelian).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	etymology	Etymology. The specific name honors Alain Cluzaud (Pessac, France), who collected part of the material here presented and greatly contributed to the knowledge of the malacofauna of the Aquitaine Basin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	materials_examined	Material examined. Lower Oligocene: France: Gaas: type material plus 66 valves (AC, BD 529, DA, JVC, Fig. 64 G, PR). Lower Miocene: France (Burdigalian): Carrière Vives: 3 valves (BD 530, JFL, Fig. 64 H). Maximum width of the valves: 3.6 / 6.8 / 5.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, moderately elevated, rounded, without trace of apex, tail valve semicircular, mucro subcentral. Tegmentum sculptured with irregularly disposed granules, elevated and closely spaced in HV, LA, PMA, loosely longitudinally arranged, some granules joined together in CA, AMA, tegmentum fully covered by aesthetes, both on granules and between them. Articulamentum with apophyses rounded, slit formula: 11 / 2 / 9 – 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, slope straight. Intermediate valves broadly rectangular (W / L = 2.75 – 2.92), moderately elevated (H / W = 0.39 – 0.42), rounded in anterior profile, anterior margin almost straight on jugal side, side margins slightly rounded, posterior margin straight, without trace of apex, lateral areas poorly elevated. Tail valve semicircular (W / L = 1.46 – 1.78), anterior margin almost straight, mucro subcentral, antemucronal slope slightly convex, postmucronal slope slightly concave just under mucro. HV, LA and PMA sculptured with closely spaced, elevated, irregularly distributed and irregularly shaped granules. CA and AMA sculptured with irregularly disposed granules, loosely longitudinally arranged, some granules joined together, others splitting in various areas, mainly near anterior margin and towards LA, more irregular in JA. Tegmentum fully covered by aesthetes, both on granules (a subcentral megalaesthete and up to 8 – 10 micraesthetes in PMA) and between them. Articulamentum with apophyses rounded, jugal sinus wide, insertion lamina divided into rather large, irregular teeth, slit formula: 11 / 2 / 9 – 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	discussion	Remarks. The fossil record of Ischnochiton cluzaudi sp. nov. is limited to the lower Oligocene and lower Miocene of the Aquitaine Basin (France). A lot of valves from the lower Oligocene and lower Miocene of the Aquitaine Basin has been attributed to Ischnochiton korytnicensis Bałuk, 1971 by Dell’Angelo et al. (2018 a, 2020 b), and really the two species (I. korytnicensis and I. cluzaudi sp. nov.) are at first glance similar. Ischnochiton cluzaudi sp. nov. shows CA and AMA sculptured with granules irregularly longitudinally arranged, very different by the quite regular longitudinal and parallel ribs present in I. korytnicensis. This difference in the sculpture of CA and AMA is sufficient to separate the valves of I. cluzaudi sp. nov. from those of I. korytnicensis (compare Figs 70 C and 64 D). We illustrate a valve showing coalescence of the last two plates into one plate vii-viii (Fig. 64 G), rarely described in fossil valves [e. g., Dell’Angelo & Forli 1995 b for two intermediate valves of Rhyssoplax saeniensis Laghi, 1984 from the Pliocene of Serre di Rapolano (Siena, Italy). Comparisons. See Tabs 9, 10 for a comparison with the Ischnochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF664E9A0FADFF0568FD9698.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b; this study). Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Carrière Vives (Dell’Angelo et al. 2018 a; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	description	Fig. 65	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	materials_examined	Type material. Holotype, MSNG 62644, tail valve, width 3.5 mm (Figs 65 I – L). Paratype 1, MSNG 62645, intermediate valve, width 3.3 mm (Figs 65 F – H). Paratype 2, MSNG 62646, head valve, width 3.7 mm (Figs 65 A – B). Paratype 3, MNHN. F. A 98472, intermediate valve, width 5 mm (Figs 65 C – E) from Maureilhan (France). Paratype 4, MHNBx 2024.3.1, intermediate valve, width 4.3 mm, from Coupe du fossé près de La Solitude (France). Type locality. Noaillan, Gamachot (France). Type stage. Lower Miocene (Burdigalian).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	etymology	Etymology. The name honors Jean-François Lesport (Sainte-Hélène, France), who collected part of the material here discussed. for his contribution to the knowledge of the malacofauna of the Aquitaine Basin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	materials_examined	Material examined. Lower Miocene (Burdigalian): France: type material plus Carrière Vives: 12 valves (JFL, PR), Coupe du fossé près de La Solitude: 1 valve (JFL), Gamachot: 63 valves (AC, BD 531, JFL, PR), Lahitet: 1 valve (PR), Maureilhan: 8 valves (BD 532). Middle Miocene (Serravallian): France: Carré: 1 valve (DA). Maximum width of the valves: 7.5 / 6.2 / 4.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, elevated, rounded, without trace of apex, tail valve semicircular, mucro subcentral to slighty anterior, swollen but not prominent. Tegmentum sculptured with concentric, large and irregular ribs in HV, LA, PMA, ribs continuing longitudinally in CA, AMA, someone joined together, others splitting, tegmentum fully covered by aesthetes, both on nodular elevations and between them. Articulamentum with apophyses small, slit formula: 12 – 13 / 2 / 12 – 14.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.35 – 3.00), moderately to highly elevated (H / W = 0.40 – 0.51), regularly rounded in anterior profile, anterior margin straight on jugal side, side margins slightly rounded, posterior margin straight, without trace of apex, lateral areas poorly elevated. Tail valve semicircular (W / L = 1.50 – 1.76), mucro subcentral to slighty anterior, swollen but not prominent, antemucronal slope almost straight, postmucronal slope almost straight or slightly concave just under mucro. HV, LA and PMA sculptured with concentric, large and irregular ribs, very close to each other; ribs continue longitudinally on CA and AMA, more irregular, someone joined together, others splitting in various areas, mainly near anterior margin and towards LA. Tegmentum fully covered by megalaesthetes and micraesthetes, both on nodular elevations and between them. Articulamentum with apophyses small, rounded, trapezoidal in tail valve, jugal sinus wide, insertion lamina divided into irregular teeth, slit formula: 12 – 13 / 2 / 12 – 14.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	discussion	Remarks. The fossil record of Ischnochiton lesporti sp. nov. is limited to the lower and Middle Miocene of the Aquitaine Basin (France). A lot of valves from the Burdigalian and Serravallian of the Aquitaine Basin has been attributed to Ischnochiton rissoi (Payraudeau, 1826) by Dell’Angelo et al. (2018 a), and really the two species (I. rissoi and I. lesporti sp. nov.) are very similar. Ischnochiton lesporti sp. nov. shows LA sculptured with concentric, almost longitudinal ribs, which continue longitudinally in CA, arranged more irregularly than in I. rissoi. This difference in the sculpture is considered sufficient to warrant separation of I. lesporti sp. nov. from I. rissoi. Comparisons. The species is close to Ischnochiton sigwartae sp. nov. (see above and Tab. 9). Ischnochiton lesporti sp. nov. is superficially similar to I. striolatus (Gray, 1828), a living species from the western Atlantic frontage (from North Carolina to Brazil); it differs from I. lesporti sp. nov. mainly by the different sculptures of CA and AMA and the geographic / stratigraphic distribution (Kaas & Van Belle 1990).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF654E9B0FADFC646A919298.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Carrière Vives, Coupe du fossé près de La Solitude, Gamachot, Lahitet, Maureilhan (Dell’Angelo et al. 2018 a; this study). Middle Miocene: northeastern Atlantic (Serravallian): Aquitaine Basin, France: Carré (Dell’Angelo et al. 2018 a; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF634E9D0FADFF056BCF95E8.taxon	description	Fig. 66	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF634E9D0FADFF056BCF95E8.taxon	materials_examined	Type material. Holotype RGM 1364007, intermediate valve, width 3.4 mm (Figs 66 A – C). Paratype RGM 1364008, intermediate valve (Figs 66 D – E). Type locality. Vale de Freixo site, near the village of Carnide, Pombal municipality (central-west Portugal). Type stage. Pliocene (upper Zanclean to lower Piacenzian), Carnide Sandstone Formation. Material examined. Pliocene: Portugal: Vale de Freixo: type material plus one intermediate valve (width 3.8 mm, valve lost, Figs 66 F – H).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF634E9D0FADFF056BCF95E8.taxon	description	Description. Head and tail valves unknown. Intermediate valves broadly rectangular, width four times length or a little more (W / L = 4.00 – 4.10), slightly elevated (H / W = 0.20 – 0.24), carinate in anterior profile, anterior margin straight, side margins rounded, posterior margin straight on both sides of prominent apex, lateral areas poorly elevated, sharply delimited, Tegmentum with microgranulose sculpture. LA sculptured with 3 radiating irregular granulose ribs originating at apex, some ribs splitting towards anterior margin, with scattered elevated granules irregularly distributed on striae. CA sculptured with longitudinal irregular granulose ribs (9 – 11 on each side), with granules coalescent forming uninterrupted ribs, interstices smooth, equal in width to ribs, ribs not reaching anterior margin near JA, large smooth triangular JA, weak growth lines. Articulamentum with large, triangular apophyses, insertion plate short with a single slit, slit rays scarcely visible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF634E9D0FADFF056BCF95E8.taxon	discussion	Remarks. Ischnochiton loureiroi Dell’Angelo, Landau, Silva & Sosso, 2022 is only known from the original material, based on a few and poorly preserved intermediate valves from the Pliocene of Portugal, but the sculpture of the valves is different from that of any other Ischnochiton spp. recorded from the European Neogene. Comparisons. The only other species with a carinated profile, among the Ischnochiton spp. with granulose ribs from the NE Atlantic area discussed here, are I. zbyi Dell’Angelo & Silva, 2003 and I. renardi Dell’Angelo, Lesport, Cluzaud & Sosso, 2018. Ischnochiton loureiroi markedly differs from I. zbyi by the shape and the different sculpture of the intermediate valves, and from I. renardi by the different sculpture of CA and LA (see Tab. 9).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF634E9D0FADFF056BCF95E8.taxon	distribution	Distribution. Lower to upper Pliocene: northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo et al. 2022).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF624E9E0FADFEB468909604.taxon	description	Fig. 67	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF624E9E0FADFEB468909604.taxon	materials_examined	Type material. Holotype MNHN. F. A 67095, tail valve, width 6.3 mm (Figs 67 A – C). Paratypes: MNHN. F. A 67096 – A 67101 (6 valves, Figs 67 D – I); NHMW 2017 / 0108 / 0020 – 0025 (6 valves, Figs 67 J – L); RGM. 1008375 – 1008380 (6 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): France: Saint-Clément-de-la-Place: type material plus 635 valves (MNHN. F. A 67102, NHMW 2017 / 0108 / 0026, RGM. 1008352, RGM. 1008381, RGM. 1008432, BD 141; Sceaux d’Anjou, La Presselière: 5 valves (BD 142). Maximum width of the valves: 6.5 / 9 / 7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF624E9E0FADFEB468909604.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight. Intermediate valve broadly rectangular (W / L = 2.25 – 2.28), rounded in anterior profile, moderately elevated (H / W = 0.37 – 0.42), anterior and posterior margins almost straight, side margins rounded, apex hardly or not developed, lateral areas hardly raised. Tail valve semicircular (W / L = 1.42 – 1.60), anterior margin slightly convex, mucro central, not prominent, antemucronal slope straight or slightly convex, postmucronal slope almost straight or slightly concave. Tegmentum smooth and glossy. HV, LA and PMA smooth with several concentric growth lines. CA and AMA smooth, with some concentric growth lines, continuing from LA and PMA, though less strongly, in some cases with presence of hardly perceptible, short longitudinal ridges in area near anterior margin. Articulamentum with apophyses rounded, separated by wide jugal sinus, slit formula: 11 / 1 / 9 – 13, slits inequidistant, slit rays clearly visible, on intermediate valves (and even less accentuated on tail valves) there are no slit rays formed by individual holes, but characteristic band of rays, teeth short, upper side roughened.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF624E9E0FADFEB468909604.taxon	discussion	Remarks. The fossil record of Ischnochiton nitidum Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is limited to the upper Miocene of the Ligerian Basin, France. The valves are large, but generally not well preserved; complete valves are scarce in the studied material. A narrower band of rays is sometimes visible in the articulamentum near the posterior margin of intermediate valves, giving the impression of the presence of a second slit in the insertion plate. Comparisons. The smooth and glossy tegmentum of Ischnochiton nitidum is characteristic, and different from that of other extant species of Ischnochiton from the Mediterranean and east Atlantic, which are characterized by a more or less marked longitudinal and radial tegmental sculpture (e. g., the Ischnochiton spp. discussed here, see tab. 7, and I. obtusus Carpenter in Pilsbry, 1893 from Portugal) or by a microgranulose sculpture [e. g., I. cessaci (de Rochebrune, 1881), I. goreensis Thiele, 1909 and I. nicklesi Kaas & Van Belle, 1990, all from the West Africa]. Ischnochiton nitidum is superficially similar to the living arctic-boreal species Stenosemus albus (Linnaeus, 1767) (see below), from which it differs by the tegmentum (uniformly microgranulose in S. albus), the different shape of the tail valve, the profile of intermediate valves (subcarinate in S. albus) and the width of the apophyses (greatly expanded, almost coalescing across the small jugal sinus in S. albus).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF624E9E0FADFEB468909604.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place, Sceaux d’Anjou Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF614E9F0FADFC186B649724.taxon	description	Fig. 68	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF614E9F0FADFC186B649724.taxon	materials_examined	Type material. Holotype: MHNBx 2017.8.1, an intermediate valve, width 6.4 mm (Figs 68 A – C). Paratypes: MZB 32135, an intermediate valve; MHNBx 2017.8.2, an intermediate valve (Fig. 68 D). Type locality. Moulin-Pochas, Ligerian Basin (France). Type stage. Upper Miocene (Messinian?). Material examined. Upper Miocene: France (Messinian?): Moulin-Pochas (Amberre): type material plus 4 valves (PR), maximum width of the valves: 7.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF614E9F0FADFC186B649724.taxon	description	Description. Head and tail valves unknown. Intermediate valves broadly rectangular, carinated in anterior profile, moderately elevated (H / W = 0.40), posterior margin almost straight, side margins rounded, apex inconspicuous, lateral areas slightly raised. CA sculptured with 52 – 56 longitudinal granulose riblets, tending to be more irregular towards side margins, with scarce evidence of interstices, and cover also JA. LA sculptured with 6 – 7 radial granulose riblets, larger than the longitudinal ones in CA, tending to split near side margins, with evidence of growth lines and with granules tending to coalesce near the anterior margin. Articulamentum with apophyses wide, one slit for each side.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF614E9F0FADFC186B649724.taxon	discussion	Remarks. Ischnochiton renardi Dell’Angelo, Lesport, Cluzaud & Sosso, 2018 is only know from the original material, based on 7 intermediate valves from the upper Miocene of France, sometimes incomplete and poorly preserved, especially the characters of the articulamentum are scarcely visible Comparisons. Ischnochiton renardi at a first look resembles I. zbyi Dell’Angelo & Silva, 2003, with whom it shares the carinated profile, but differs by the different sculpture, longitudinal sulci with interstices almost equal or slightly narrower than sulci and smooth jugum in I. zbyi, granulose riblets with scarce evidence of interstices, and covering also the jugal area in I. renardi, and the greatest elevation (H / W = 0.40 vs. 0.26 – 0.36 in I. zbyi), and moreover the different geographic and stratigraphic range, upper Miocene of Ligerian Basin (France) for I. renardi, vs. Pliocene of Mondego Basin (Portugal) for I. zbyi.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF614E9F0FADFC186B649724.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic: Ligerian Basin, France: Moulin Pochas (Dell’Angelo et al. 2018 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF604E800FADFDF86B8D9420.taxon	description	Fig. 69	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF604E800FADFDF86B8D9420.taxon	materials_examined	Type material. Holotype: MNHN / UL. II. 406 (intermediate valve). Paratypes: MNHN / UL. II. 407 - 408 (head and tail valves); MZB 12692 (3 valves); BD FW 49 (3 valves); GeoFCUL VFX. 03.312 - 332 (21 valves). Type locality. Vale de Freixo site, near the village of Carnide, Pombal municipality (central-west Portugal). Type stage. Pliocene (upper Zanclean to lower Piacenzian), Carnide Sandstone Formation. Material examined. Pliocene: Portugal: Vale de Freixo: type material plus 505 valves (GeoFCUL VFX. 03.334 – 336, GeoFCUL VFX. 03.351 – 354, RGM. 1364002 – 1364006, MNHN. F. A 81983 – A 81985, BD 240, Figs 69 A – L). Maximum width of the valves: 5.8 / 6.5 / 4.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF604E800FADFDF86B8D9420.taxon	description	Description. Head valve semi-circular, posterior margin widely V-shaped, notched in middle, slope straight to slightly concave. Intermediate valves broadly rectangular, three times wider than long or more (W / L = 3.00 – 3.50), moderately elevated (H / W = 0.26 – 0.36), carinate in anterior profile, anterior margin straight, slightly concave in jugum, side margins straight to slightly rounded, posterior margin straight, with apex not evident, lateral areas moderately elevated, jugal area triangular. Tail valve semicircular, depressed (W / L = 1.90 – 2.10), anterior margin straight or slightly concave in jugal tract, mucro subcentral, slightly raised, antemucronal slope straight, postmucronal slope slightly concave directly behind mucro. Tegmentum rough. HV, LA and PMA sculptured with microgranulose sculpture of roundish / squarish granules up to 30 – 40 μm, with 3 – 5 aesthetes of equal size, and granulose radiating ribs (HV 19 – 24, not starting from apex, but from mid-valve, concentrically crossed by numerous growth lines, some ribs splitting towards anterior margin; LA 2 – 8, starting from apex, with microgranules always present between ribs; PMA 24 – 30), granules larger towards margins (up to 100 μm, with 10 – 15 or more aesthetes of equal size), with microgranules present in all rib interspaces. CA and AMA sculptured with longitudinal granulose ribs (10 – 26 on either side), with granules coalescent forming uninterrupted ribs, interstices narrower than ribs with subobsolete microgranulose sculpture, JA triangular with longitudinal striae converging towards center on both sides in wider valves, forming larger, irregular granules; not arranged in longitudinal striae in smaller valves. Articulamentum whitish, apophyses wide, triangular in intermediate valves, trapezoidal in tail valve, jugal sinus narrow, slit formula 10 – 11 / 1 / 7 – 11, slits inequidistant.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF604E800FADFDF86B8D9420.taxon	discussion	Remarks. The fossil record of Ischnochiton zbyi Dell’Angelo & Silva, 2003 seems limited to the Pliocene of Portugal. The abundant material studied, representing the complete growth series from juvenile to adult, has permitted a better understanding of intraspecific variability (Dell’Angelo et al. 2022), e. g., the sculpture of intermediate valves, with the granulose radial ribs in the lateral areas ranging from 5 – 8 in the larger valves (Fig. 69 D) to 2 – 3 in smaller valves (Fig. 69 J), with microgranules always present in the rib interspaces, and the number of longitudinal ribs in the pleural areas varying from 22 – 26 on either side in the larger valves (Fig. 69 D) to 10 – 15 in juveniles (Fig. 69 J). Comparisons. This species is similar to Ischnochiton crovatoi Dell’Angelo, Sosso & Bonfitto, 2019, from which it differs by the shape of the tail valve, more squared in I. crovatoi (W / L = 1.71) respect to I. zbyi (W / L = 1.90 – 2.10), the greater number of radial riblets on the postmucronal area (42 in I. crovatoi vs. 24 – 26 in I. zbyi) and, mainly, by the absence in I. crovatoi of any concentric grooves on HV, LA and PMA, concentric grooves instead well present in I. zbyi.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF604E800FADFDF86B8D9420.taxon	distribution	Distribution. Pliocene: northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo & Silva 2003; Dell’Angelo et al. 2022).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7F4E820FADFAFA6B9C97F0.taxon	description	Fig. 70	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7F4E820FADFAFA6B9C97F0.taxon	description	Dell’Angelo 1998, p. 78; Dulai 2005, p. 34; Dulai & Studencka 2007, p. 17; Studencka & Dulai 2010, p. 265; Dell’Angelo et al. 2013, p. 77, pl. 3, figs L – T; Dell’Angelo et al. 2015 a, p. 234, pl. 5, figs 1 – 9; Ruman & Hudácková 2015, p. 166; Dell’Angelo et al. 2016, p. 96; Dell’Angelo et al. 2018 b, p. 53, tab. 17; Dell’Angelo et al. 2021 b, p. 413, figs 54 – 61. Ischnochiton (Simplischnochiton) korytnicensis; Dell’Angelo et al. 1999, p. 267, pl. 3, figs 1 – 2. non Ischnochiton cf. korytnicensis; Studencka & Studencki 1988, p. 45 (= I. rissoi, fide Studencka & Dulai 2010: 265). non Ischnochiton korytnicensis; Dell’Angelo et al. 2018 a, p. 23, figs 5 J – L; Dell’Angelo et al. 2020 b, p. 5, fig. 2 (= I. cluzaudi sp. nov., fide this study) Ischnochiton rudolticensis (non Ischnochiton rudolticensis Šulc, 1934); Bałuk 1965, p. 369, pl. 1, fig. 7 (fide Bałuk 1984). Ischnochiton rissoi (non Ischnochiton rissoi Payraudeau, 1826); Laghi 1977, p. 104, pl. 1, fig. 9 (partim, non figs 4 – 8 = I. rissoi).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7F4E820FADFAFA6B9C97F0.taxon	materials_examined	Type material. Holotype in Bałuk’s collection, reg. No. BkK-A 15, an intermediate valve illustrated in Bałuk (1971: pl. 3, fig. 2). Type locality. Korytnica, southern slopes of the Holy Cross Mts. (Poland). Type stage. Middle Miocene. Material examined. Middle Miocene: Central Paratethys: Romania: Lapugy: 1 valve (NHMW 2010 / 0256 / 0005); Eastern Paratethys: Ukraine: Varovtsi: 21 valves (BD 533, Figs 70 E – J). Miocene (Tortonian): Italy: Borelli: 9 valves (PG, MZB 32026 – 32027, MGPT PU 135041), Montegibbio: 21 valves (BD 534, MZB 32021 – 32022, Figs 70 C – D), Rio di Bocca d’Asino: 39 valves (BD 535, Figs 70 K – L, PG, MZB 32023 – 32025), Villa Monti: 3 valves (BD 536, MZB 32050). Lower Pliocene: Italy: Liguria: Borzoli: 2 valves (BD 537, MZB 45712); Bussana: 15 valves (BD 538, PG, MZB 45714 – 45715), Genova Sestri: 1 valve (MZB 45710), Rio Sant’Antonino: 3 valves (MP), Rio Torsero: 2 valves (BD 539, MZB 45711, MZB 45713). Pliocene: Italy: Piedmont: Vintebbio: 26 valves (BD 540, Figs 70 A – B); Emilia-Romagna: Cava di Campore: 1 valve (BD 541). Maximum width of the valves: 8.3 / 9.5 / 7.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7F4E820FADFAFA6B9C97F0.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.56 – 2.72), moderately elevated (H / W = 0.44), rounded in anterior profile, anterior margin almost straight on jugal side, side margins slightly rounded, posterior margin straight, without trace of apex, lateral areas moderately elevated. Tail valve semicircular (W / L = 1.50 – 1.79), anterior margin almost straight, mucro subcentral, antemucronal slope slightly convex, postmucronal slope slightly concave just under mucro. HV, LA and PMA sculptured with closely spaced, elevated, irregularly distributed and irregularly shaped nodules, which in some cases, especially in PMA, give the impression to be arranged in radial rows. CA and AMA sculptured with quite regular longitudinal ribs, more spaced near lateral margins, more irregular in JA. Tegmentum fully covered by aesthetes, both on nodular elevations (without a clearly recognizable structure of megalaesthetes and micraesthetes) and between them. Articulamentum with apophyses rounded, jugal sinus wide, insertion lamina divided into rather large, irregular teeth, slit formula: 11 – 12 / 1 / 12.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7F4E820FADFAFA6B9C97F0.taxon	discussion	Remarks. This species was described from the Middle Miocene (Badenian) of Poland, and was subsequently found in several other deposits, from the Miocene to Pliocene, which extended its geographic and stratigraphic distribution outside the Paratethys. Ischnochiton korytnicensis Bałuk, 1971 is similar to I. rissoi (Payraudeau, 1826), and the two species can coexist in the same sites, e. g., for the Miocene of Korytnica (Bałuk 1971, 1984) and N. Italy (Dell’Angelo et al. 2015 a), and for the Pliocene of Liguria (Dell’Angelo et al. 2013). Comparisons. This species is similar to Ischnochiton rissoi (Payraudeau, 1826), from which it differs mainly in the ornamentation of HV, LA and PMA composed of irregular, closely spaced, and elevated nodules, whereas I. rissoi shows concentric vermicular or more or less radial ribs.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7F4E820FADFAFA6B9C97F0.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Poland: Korytnica, Łychów, Niskowa (Bałuk 1965, 1971, 1984; Macioszczyk 1988), Romania: Lapugy (this study); Eastern Paratethys: Ukraine: Varovtsi (this study); Upper Miocene: Proto-Mediterranean Sea (Tortonian): Italy: Borelli, Montegibbio, Rio di Bocca d’Asino, Villa Monti (Laghi, 1977; Dell’Angelo et al. 1999, 2015 a). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Genova Sestri, Rio Sant’Antonino, Rio Torsero (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b). Pliocene: central Mediterranean, Italy: Piedmont: Vintebbio; Emilia-Romagna: Cava di Campore (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	description	Fig. 71	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	materials_examined	Type material. Holotype: MSNG 62647, intermediate valve, width 4.8 mm (Figs 71 B – D). Paratype 1: MSNG 62648, head valve, width 3.5 mm (Fig. 71 A). Paratype 2: MSNG 62649, tail valve, width 4 mm (Figs 71 I – L). Paratype 3: MNHN. F. A 98473, intermediate valve, width 3 mm. Paratype 4: NHMW xxx, intermediate valve, width 3.3 mm (Figs 71 E – G). Paratype 5: NHMW 2010 / 0256 / 0005 (as I. rudolticensis), intermediate valve, width 2.7 mm (Figs 71 M – P) from Lapugy (Romania). Type locality. Varovtsi (Ukraine). Type stage. Middle Miocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	etymology	Etymology. The name honors Andreas Kroh (NHMW), in recognition of his many paleontological contributions on European Cenozoic marine invertebrates.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	materials_examined	Material examined. Middle Miocene: Eastern Paratethys: Ukraine: Varovtsi: type material plus 5 valves (BD 252, Fig. 71 H, BD 253). Maximum width of the valves: 3.5 / 4.8 / 4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, width about three times length, moderately elevated, rounded, without trace of apex, tail valve semi-circular, mucro subcentral, not prominent. Tegmentum sculptured with large and irregular granules, seeming radial striae in HV, LA, PMA, forming a network in CA, AMA, tegmentum fully covered by aesthetes, both on nodular elevations and between them. Articulamentum with apophyses rounded, slit formula: - / 2 / 11.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular, width about three times length (W / L = 2.82 – 3.01), moderately elevated (H / W = 0.34), rounded in anterior profile, anterior margin almost straight, side margins slightly rounded, posterior margin straight, without trace of apex, lateral areas not elevated. Tail valve semicircular, W / L = 1.68 – 1.82, anterior margin straight, mucro subcentral, not prominent, antemucronal slope straight to slightly convex, postmucronal slope slightly concave. HV, LA and PMA sculptured with large and irregular granules, very close to each other, randomly oriented, aligned along growth lines towards margins, but giving the impression to form radial striae of granules, especially in HV and PMA. CA and AMA sculptured with irregular elongated granules, slightly overlapping each other, less evident in JA, arranged randomly but giving the impression of forming a network. Tegmentum fully covered by aesthetes, both on nodular elevations (without a clearly recognizable structure of megalaesthetes and micraesthetes) and between them. Articulamentum with apophyses rounded, trapezoidal in tail valve, jugal sinus wide, insertion lamina divided into irregular teeth, slit formula: - / 2 / 11.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	discussion	Remarks. The fossil record of Ischnochiton krohi sp. nov. is limited to the Middle Miocene (Badenian) of Paratethys (Romania and Ukraine). The material is scarce, a few valves but quite well preserved. A tail valve (Fig. 71 H) shows a slightly different shape, more circular that those of the other tail valves examined, decidedly semicircular (e. g., Fig. 71 I), but we consider that it can be included in the intraspecific variability of Ischnochiton krohi sp. nov. Comparisons. The more similar species is Ischnochiton ligusticus Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013, from which I. krohi sp. nov. differs mainly by the different sculpture, consisting of very irregular granules, arranged in segments of various size and shape, slightly overlapping each other, forming rugosities in I. ligusticus, while the granules are more regular in I. krohi sp. nov., giving the impression to be arranged in radial striae in HV and PMA. Also, the stratigraphic distribution is different (Middle Miocene of Paratethys for I. krohi sp. nov. vs. upper Miocene to lower Pliocene of Italy for I. ligusticus), and the number of slits in intermediate valves, 2 in I. krohi sp. nov. vs. 1 in I. ligusticus.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Romania: Lapugy; Eastern Paratethys: Ukraine: Varovtsi (this study). Genus Stenoplax Carpenter MS, Dall, 1879	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	type_taxon	Type species. Chiton limaciformis Sowerby in Broderip & Sowerby, 1832, by original designation.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	distribution	Distribution. Stenoplax is known from the Eocene to the Recent, with a living distribution mostly in temperate or tropical regions of the central and eastern Pacific (Hawaii, and from Canada to Peru), Indo – Pacific (from Japan to India), Indian Ocean (Madagascar), East Atlantic Ocean (Gulf of Guinea), West Atlantic Ocean (from Florida to Brazil including the Caribbean) (Kaas & Van Belle 1988). The fossil record extends back to the middle Eocene in Europe [U. K. (Wrigley 1943; Cherns & Schwabe 2019) and Ukraine (Bielokrys 1999)] and the upper Eocene or lower Oligocene in Washington, U. S. A. (Puchalski et al. 2008; Dell’Angelo et al. 2011 a), the lower Oligocene in Italy (Dell’Angelo & Palazzi 1992; Dell’Angelo et al. 2015 a), the Miocene in Italy (Dell’Angelo et al. 2014) and France (Dell’Angelo et al. 2018 a), and the Pliocene-Pleistocene of California, U. S. A. (Berry 1922; Vendrasco et al. 2012 and the references listed therein) and Japan (Itoigawa et al. 1976, 1978).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7D4E840FADFC8C6FE59282.taxon	discussion	Remarks. Stenoplax is a distinctive genus whose members are markedly elongate and whose intermediate valves have prominent sutural laminae and generally raised lateral areas. The depressed and much more elongated (relative to other valves) tail valve is diagnostic for this genus. The main morphological characters of the Stenoplax spp. considered in the present study are reported in Tab. 11, together with the species of the genus Connexochiton Kaas, 1979. ...... continued on the next page	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7A4E870FADF946682D9794.taxon	description	Fig. 72	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7A4E870FADF946682D9794.taxon	materials_examined	Type material. Holotype: MGPT PU 108791 (tail valve). Paratypes: MGPT PU 108792 – 108795: 4 valves (Fig. 72 J); MSNG 57252: 2 valves; ZISP 2199 – 2200: 2 valves; MZB 60085 – 60087: 3 valves (Figs 72 I, 72 K – L); NHMW 2013 / 0272 / 0001: 2 valves. Type locality. Rio di Bocca d’Asino, Alessandria (Italy). Type stage. Miocene (Tortonian). Material examined. Middle Miocene (Langhian): Italy: Albugnano: 1 valve (MZB 32030). Upper Miocene (Tortonian): Italy: Borelli: 1 valve (MGPT PU 135042), Rio di Bocca d’Asino: type material plus 37 valves (BD 542, PG, MZB 32031 – 32032), Villa Monti: 1 valve (MZB 32052). France: Beugnon: 8 valves (RGM. 1310172, RGM. 1310175, RGM. 1310179), Renauleau: 5 valves (BD 144, MNHN. F. A 67107), Saint-Clément-de-la-Place: 347 valves (BD 143, MNHN. F. A 67103 – A 67106, Figs. 72 A – H, NHMW 2017 / 0108 / 0027, RGM. 1008353, RGM. 1008382, RGM. 1008433). Maximum width of the valves: 7 / 8.8 / 5.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7A4E870FADF946682D9794.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.27), moderately elevated (H / W = 0.38), rounded in anterior profile, anterior margin straight between apophyses, side margins rounded, posterior margin about straight, apex not indicated, lateral areas little raised, poorly defined. Tail valve semicircular, depressed (W / L = 1.52 – 1.66), anterior margin straight, mucro subcentral, weak indicated, antemucronal and postmucronal slopes almost straight. HV, LA and PMA sculptured with many close-set, irregularly undulate concentric grooves, better defined towards periphery. CA and AMA smooth, with some longitudinal grooves (that continue without interruption concentric grooves) towards the margins, slightly obliquely directed. Articulamentum with apophyses triangular, widely separated by straight sinus, insertion plates short, slit formula 10 – 13 / 2 / 14, slit rays neatly indicated by distinct rays of pores, teeth sharp, eaves narrow.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7A4E870FADF946682D9794.taxon	discussion	Remarks. Stenoplax paviai Dell’Angelo, Giuntelli, Sosso & Zunino, 2014 was originally described on a few valves from the Miocene (Tortonian) of Rio di Bocca d’Asino, complete intermediate valves were not present, and some important characters were unknown. New and abundant material found in recent years from the Miocene (Tortonian) of Saint-Clément-de-la-Place allowed to complete the description also for the intermediate valves and to better define some specific characters (Dell’Angelo et al. 2018 b). The fossil record was also extended to the Miocene (Langhian) of N. Italy (Dell’Angelo et al. 2015 a). Comparisons. Stenoplax paviai Dell’Angelo, Giuntelli, Sosso & Zunino, 2014 differs from S. veneta Dell’Angelo & Palazzi, 1992 by the different sculpture, fully covered by small granules in S. veneta. Stenoplax paviai is superficially like Tegulaplax hululensis (E. A. Smith, 1903), a living species with a widespread distribution in the Indo-Pacific, and also in the Mediterranean coast of Israel, from which it differs by the different shape of the valves, the sculpture CA and AMA, the pectinate teeth (Kaas et al. 2006).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF7A4E870FADF946682D9794.taxon	distribution	Distribution. Lower Miocene: Proto – Mediterranean Sea (Langhian): N. Italy: Albugnano (Dell’Angelo et al. 2015 a). Upper Miocene: northeastern Atlantic: Ligerian Basin, France: Beugnon, Renauleau, Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b); Proto – Mediterranean Sea: Po Basin, N. Italy: Borelli, Rio di Bocca d’Asino, Villa Monti (Dell’Angelo et al. 2014, 2015 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	description	Fig. 73	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	materials_examined	Type material. Holotype MZB 10301, tail valve. Paratype BD 3039, a head and 2 tail valves (Figs 73 A – D). Type locality. Case Soghe, Vicenza (Veneto, Italy). Type stage. Lower Oligocene (Rupelian). Material examined. Lower Oligocene: Italy: Case Soghe: type material plus 5 valves (2 intermediate and 3 tail, BD 543, Figs 73 E – H). Maximum width of the valves: 2.8 / 3 / 4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	description	Description. Head valve probably semicircular (only a fragment known). Intermediate valve broadly rectangular, anterior margin straight, posterior margin seems straight for the visible part, lateral area poorly defined, not distinctly separated by central area, only a sign of diagonal fold between lateral and central areas is present but hardly evidenced. Tail valve semielliptical (W / L = 1.24), depressed, anterior margin almost straight, mucro subcentral, not elevated, antemucronal slope straight or slightly convex, postmucronal slope almost straight just behind mucro. Tegmentum fully covered by small granules, irregularly disposed and some coalescing in HV, LA and PMA, with marked concentric lines of growth visible on PMA, arranged in groups that seem obliquely directed in CA and AMA. Articulamentum with trapezoidal apophyses in tail valve, separated by a straight sinus, teeth short and irregular, slit formula:? / 1? / 12 – 16.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	discussion	Remarks. This is the sole described polyplacophoran species from the Oligocene of Italy. Originally it was described on four valves only (1 head fragment and 3 tail), and the morphology of the intermediate valves was unknown. New material found in recent years (2 intermediate and 3 tail valves) allowed to complete the description also for the intermediate valves and to better define some specific characters (Dell’Angelo et al. 2015 b). Not all characters of the species match those of the genus Stenoplax Carpenter MS, Dall, 1879. The lateral areas of Stenoplax spp. are generally rather elevated (Kaas & Van Belle 1987: 124; Vendrasco et al. 2012: 34) but are little raised and poorly defined in S. veneta Dell’Angelo & Palazzi, 1992. However, the same characteristic is seen in other Stenoplax species, e. g., S. marcusi (Righi, 1971). Comparisons. The tegmentum fully covered by small granules of Stenoplax veneta is characteristic, and different from that of S. paviai Dell’Angelo, Giuntelli, Sosso & Zunino, 2014.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	distribution	Distribution. Oligocene (Rupelian): N. Italy: Case Soghe (Dell’Angelo & Palazzi 1992; Dell’Angelo et al. 2015 b). Genus Stenosemus von Middendorff, 1847	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	type_taxon	Type species. Chiton albus Linnaeus, 1767, by subsequent designation (Winckworth 1926).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	distribution	Distribution. The genus Stenosemus is known from the Miocene up to Recent. Stenosemus is the most widely distributed genus of the order Chitonida, with as many as 24 living species described thus far, from the Arctic to Antarctic regions, including cold deep waters of tropical region, mainly at depths greater than 200 m down to 4572 m (Sirenko 2016, 2017). Its fossil record includes the lower Miocene of France (Dell’Angelo et al. 2020 b), the middle Miocene of Hungary (Schwabe & Dulai 2024), the upper Miocene of France and Italy (Dell’Angelo et al. 2015 a, 2018 a), the Pliocene of Spain and Italy (Dell’Angelo et al. 2004, 2012, 2013; this study), the Pleistocene of Italy (Dell’Angelo & Giusti 1997; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF784E890FADFD686D919391.taxon	discussion	Remarks. The genus Stenosemus was considered by Kaas & Van Belle (1990) as a subgenus of Ischnochiton, differing mainly on the soft parts morphology (i. e. the dorsal girdle covered with conical scales in Stenosemus, with imbricating, generally striated scales in Ischnochiton). However, the first who considered Stenosemus as a distinct genus was Ferreira (1981). Regarding the Mediterranean basin, only one species, Stenosemus dolii (Van Belle & Dell’Angelo, 1998), was considered up to now for the paleontological record of the Mediterranean basin, characterized by a great variability of valves’ morphology, especially regarding the greater or lesser graininess of the radial sculpture. In this respect, the Pliocene of Altavilla Milicia (Sicily) provided abundant material attributed to Stenosemus dolii by Dell’Angelo et al. (2012). An in-depth reconsideration of such material suggests that two undescribed species are, in fact, present which we formalize here as S. juliuspisai sp. nov. and S. praedolii sp. nov. The main morphological characteristics of the Stenosemus spp. considered in the present study are reported in Tab. 12. ...... continued on the next page	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF744E8C0FADFF056E4495E8.taxon	description	Fig. 74	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF744E8C0FADFF056E4495E8.taxon	materials_examined	Type material. The Linnean Society of London (Dodge 1952). Type locality. “ In O. Islandico ”. Material examined. No fossil material available, only descriptions and illustrations from the literature. Recent: N. Atlantic: Europe: Barents Sea, Greenland, Iceland, Norway, Denmark (BD 255, Figs 74 A – H); America: Canada (Nova Scotia), Maine (BD, ca 20 spm). Maximum width of the valves: 4 / 4.5 / 3.4 mm (measured on recent valves).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF744E8C0FADFF056E4495E8.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, weakly notched in middle, front slope straight. Intermediate valves broadly rectangular, carinate in anterior profile, rather elevated (H / W = 0.40 – 0.50), anterior margin slightly angular, side margins rounded, posterior margin about straight, apex hardly or not indicated, lateral areas scarcely raised. Tail valve semicircular, length aboul half width. mucro central, slightly elevated, not prominent, antemucronal slope slightly convex, postmucronal slope straight to weakly concave. Tegmentum uniformly microgranulose, smooth and glossy to naked eye. HV, LA and PMA with several concentrical, low growth wrinkles, continuing, though less strongly, across CA and AMA. Articulamentum with apophyses short, very wide, rounded, almost coalescing across small, shallow sinus, slit formula 13 – 14 / 1 (2) / 10 – 12, slits inequidistant, slit rays hardly or not indicated, teeth short, sharp, upper side and outer edge decidedly roughened, eaves narrow, solid.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF744E8C0FADFF056E4495E8.taxon	discussion	Remarks. The fossil record of Stenosemus albus (Linnaeus, 1767) is limited to the upper Pleistocene of Scandinavia (Norway and Sweden). Comparisons. See Tab. 12 for a comparison with the Stenosemus spp. considered in the present study. Distribution. Pleistocene: N Atlantic: Norway (Grønlie 1927; Hägg 1951; Feyling-Hanssen 1955); Sweden (Antevs 1917, 1928). Recent: Northern Atlantic Ocean, from Spitsbergen to Vigo (Spain) and near the Azores (Kaas & Van Belle 1990; Van Belle 1984; Urgorri et al. 2017); Northern Pacific Ocean, SE to San Diego, California, and SW to the Okhotsk Sea and the Sea of Japan (Dall 1921; Kaas & Van Belle 1990).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF734E8E0FADFEB56E4C9604.taxon	description	Fig. 75	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF734E8E0FADFEB56E4C9604.taxon	materials_examined	Type material. Holotype: MZB 11302, intermediate valve, width 7.5 mm. Paratypes: MZB 11303 (5 valves), IRSN IG 28523 (4 valves), MNHN (3 valves), and private collections (320 valves): BD P 151 F / 01 (90 valves, Figs 75 A – H), FG (80 valves), RA (138 valves), VB F 1003 a (12 valves). Type locality. Tyrrhenian Sea, off Civitavecchia (Italy), within a Roman amphora (Latin: dolium) at a depth of 550 m (42 ° 04 ’ 04 ” N, 11 ° 45 ’ 31 ” E). Type stage. Subfossil (“ the correct geological age of I. dolii cannot be established with any degree of certainty ”). Material examined. Lower Pliocene: Italy: Liguria: Borzoli: 15 valves (BD 544, MZB 45720). Pliocene: Spain: Estepona: 1 valve (BD 545). Italy: Piedmont: Valle Botto: 5 valves (BD 546); Emilia-Romagna: Cava di Campore: 2 valves (BD 547); Sicily: Altavilla: 110 valves (AG, AR, Fig. 75 L, BD 548, DGUP). Pleistocene: Italy: Tuscany: Capraia Island-Capo Corso - 350 / 500 m: 12 valves (BD 549, Figs 75 I – K); Emilia-Romagna: Torrente Stirone: 1 valve (BD 550); Latium: off Civitavecchia: type material; Calabria: Pecoraro: 1 valve (BD 551). Maximum width of the valves: 7 / 8 / 6.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF734E8E0FADFEB56E4C9604.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, with a rounded notch at apex. Intermediate valves broadly rectangular (W / L = 2.54 – 2.57), carinate in anterior profile, moderately elevated (H / W = 0.34 – 0.49), anterior margin consisting of three segments, two lateral ones straight or slightly concave, and one between the apophyses straight or slightly convex, side margins almost straight or slightly rounded, posterior margin practically straight with mucro not very visible, lateral areas moderately elevated. Tail valve semicircular (W / L = 1.75 – 1.88), depressed, anterior margin as in intermediate valves, mucro in slightly anterior position and little elevated, antemucronal and postmucronal slopes almost straight. Tegmentum rough. HV, LA and PMA sculptured with rather flat radial ribs (HV 17 – 26 at apex, splitting near anterior margin, LA 4 – 8 tending to become obsolete near lateral margins, PMA 18 – 20, sometimes barely conspicuous around mucro), separated by rather narrow spaces, and with well-evident and numerous growth lines, which intersect the radial ribs forming more or less elongated quadrangular granules. CA and AMA sculptured with 11 – 23 longitudinal grooves on each side, normally all reaching anterior margin with JA smooth, only in some valves 2 – 4 more internal grooves may not reach margin and converging towards jugum or become very irregular. Articulamentum with apophyses short and wide, rounded, jugal sinus not very wide, weakly convex, insertion lamina divided into short and rough denticles of irregular width, slit formula 9 – 10 / 1 / 6 – 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF734E8E0FADFEB56E4C9604.taxon	discussion	Remarks. The fossil record of Stenosemus dolii (Van Belle & Dell’Angelo, 1998) is limited to the Mediterranean Pliocene (Italy and Spain) and Pleistocene (Italy). Stenosemus dolii was described on the basis of many loose subfossil valves found in a Roman amphora off Civitavecchia, at a depth of 550 m (Van Belle & Dell’Angelo 1998); additional material pertains to a taphocoenosis from the South Ligurian Sea, between Corsica and Capraia Island, collected at a depth of 350 / 500 m (Dell’Angelo & Giusti 1997, 2000). Living specimens have been found in the Tyrrhenian Sea between 150 and 480 m (Smriglio et al. 1989); originally classified as Ischnochiton vanbellei Kaas, 1985 (Dell’Angelo & Smriglio 1999; Dell’Angelo et al. 2001 b), these specimens have been transferred to the genus Stenosemus based upon the girdle’s conic scales. Dell’Angelo et al. (2018 a) attributed tentatively to Stenosemus dolii a unique tail valve from the upper Miocene of Moulin Pochas (France). Despite the existing similarity with S. dolii, it is not possible to identify with certainty this valve, that is discussed and figured in the section dedicated to the “ Species of unclear taxonomic position ”, and it is left in open nomenclature as Stenosemus sp. An in-depth reconsideration of the abundant material from the Pliocene of Altavilla Milicia (Sicily), attributed to Stenosemus dolii by Dell’Angelo et al. (2012), suggests that two undescribed species are, in fact, present which we formalize here as S. juliuspisai sp. nov. and S. praedolii sp. nov. (see below). We consider Stenosemus dolii as a species with a radial sculpture (HV, LA, PMA) consisting of flat radial ribs intersected by well-evident and numerous growth lines, forming more or less elongated quadrangular granules, different from the real granulose sculpture characteristic of the two new species described below. Laghi (1977: pl. 2, figs 1 – 4) recorded some valves from the Pliocene (Zinola, Valle Andona and Castell’Arquato) under the name Lepidozona dorsuosa (Haddon, 1886). These valves were later considered to fall in the range of variability of Stenosemus dolii by Dell’Angelo et al. (2018 a). One of the valves illustrated by Laghi (1977: pl. 2, fig. 3 a – 3 b), from the Pliocene of Castell’Arquato, agree with the attribution to S. dolii, but the other valves figured can be attributed to Stenosemus juliuspisai sp. nov. (see below). The same partial attribution to both Stenosemus juliuspisai sp. nov. and S. praedolii sp. nov. has also been considered for the material, before attributed to S. dolii, from the upper Miocene of Northern Apennines [Dell’Angelo et al. 1999, as Ischnochiton (Simplischnochiton) exaratus (Sars, 1878); Dell’Angelo et al. 2015 a, 2016, as Stenosemus dolii], the Pliocene of Liguria, Italy (Dell’Angelo et al. 2013, 2021 b, as Stenosemus dolii), the Pliocene of Estepona, Spain [Dell’Angelo et al. 2004, as Ischnochiton (Stenosemus) aff. vanbellei], the Pleistocene of Capraia Island-Capo Corso, Italy (Dell’Angelo & Giusti 1997, 2000, as Ischnochiton exaratus). The valves show a certain variability with regard to the sculpture, e. g., the number of longitudinal grooves on each side of the central areas of intermediate valves (varying from 11 to 23), the number of flat radial ribs of the lateral areas of intermediate valves (varying from 4 to 8 or more) the jugum normally smooth (Fig. 70 J), but with a sculpture of grooves converging towards the apex visible on the upper part (Fig. 70 D) when the more internal longitudinal grooves are converging towards it (normally in valve ii). Comparisons. See Tab. 12 for a comparison with the Stenosemus spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF734E8E0FADFEB56E4C9604.taxon	distribution	Distribution. Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli (Dell’Angelo et al. 2013, 2021 b). Pliocene: western Mediterranean, Estepona Basin, Spain (Dell’Angelo et al. 2004); central Mediterranean, Italy: Altavilla Milicia (Dell’Angelo et al. 2012), Castellarquato (Laghi 1977), Valle Botto, Cava di Campore (this study). Pleistocene: central Mediterranean, Italy: off Civitavecchia (Van Belle & Dell’Angelo 1998), Capraia Island-Capo Corso, - 350 / 500 m (Dell’Angelo & Giusti 1997, 2000), Pecoraro, Torrente Stirone (this study). Recent: Mediterranean Sea, off the Latium coast and in Tuscan Archipelago, between 150 and 560 m, and in association with white coral biocoenosis (Dell’Angelo & Smriglio 1999; Dell’Angelo et al. 2001 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	description	Fig. 76	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	materials_examined	Type material. Holotype: MSNG 62650, intermediate valve, width 5.7 mm (Figs 76 A – C). Paratype 1: MSNG 62651, head valve (Figs 76 D – E). Paratype 2: MSNG 62652, tail valve (Figs 76 F – H). Paratype 3: MNHN. F. A 98474, intermediate valve, width 4.8 mm. Paratype 4: MGGC 23374, tail valve, width 3.9 mm. Paratype 5: MSNG 62653, intermediate valve, width 3.4 mm (Figs 76 J – K), from Capraia Island-Capo Corso - 350 m, Pleistocene (presumably last glacial). Type locality. Altavilla Milicia (Sicily, Italy). Type stage. Pliocene-Pleistocene (upper Piacenzian to lower Gelasian) (Dominici et al. 2020).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	etymology	Etymology. This species honors the Italian geologist and paleontologist Giulio Pisa (1936 - 1976), expert in Triassic ammonites, for his role in stimulating the malacological research of one of the Authors (MT) on the Pleistocene deposits of the Stirone river, rich in polyplacophoran remains.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	materials_examined	Material examined. Lower Pliocene: Italy: Liguria: Borzoli: 115 valves (BD 552, MZB 45723 – 45724), Bussana: 3 valves (BD 553), Rio Sant’ Antonino: 39 valves (MP, MZB 45721 – 45722), Zinola: 7 valves (BD 554). Pliocene. Spain: Estepona: 4 valves (BD 555). Italy: Piedmont: Valle Botto: 1 valve (BD 556); Emilia-Romagna: Cava di Campore: 1 valve (BD 557), Rio Merli: 2 valves (CT); Sicily: Altavilla (type material plus 25 valves, BD 558, Fig. 76 I). Pleistocene. Italy: Tuscany: Capraia Island-Capo Corso - 350 / 500 m: 7 valves (BD 559), Calabria: San Procopio: 2 valves (BD 560, Fig. 76 L). Recent. Mediterranean Sea: Spain: Alboran Island - 80 / 150 m (1 valve, BD 561, Figs 76 M – N); Italy: Alghero - 180 m (3 valves, BD 562, Figs 76 O – P). Maximum width of the valves: 4 / 5.7 / 8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, carinate in anterior profile, moderately elevated, mucro inconspicuous, tail valve semicircular, mucro subcentral and little elevated. Tegmentum rough, sculptured with radial striae of quadrangular, elevated, more or less elongated and dense nodules in HV, LA, PMA, very few splitting, and longitudinal riblets on each side in CA, AMA, JA smooth. Articulamentum with apophyses short and wide, slit formula 7 / 1 / 8 – 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, with a rounded notch at apex. Intermediate valves broadly rectangular (W / L = 2.94 – 3.26), carinate in anterior profile, moderately elevated (H / W = 0.34 – 0.48), anterior margin consisting of three segments, two lateral ones straight or slightly concave, and one between the apophyses straight or slightly convex, side margins almost straight or slightly rounded, posterior margin practically straight with mucro not very visible, lateral areas elevated. Tail valve semicircular, width about twice length (W / L = 1.90 – 2.04), anterior margin as in intermediate valves, mucro subcentral and little elevated, antemucronal slope slightly convex, postmucronal slope slightly concave. Tegmentum rough. HV, LA and PMA sculptured with radial striae of quadrangular, elevated, more or less elongated and dense nodules (HV 16 – 22 at apex, very few splitting, LA 2 – 4, one of which can split, PMA 16 – 22, a few splitting), a few growth lines, little evident. CA and AMA sculptured with 11 – 16 longitudinal riblets on each side, normally all reaching anterior margin, more irregular and converging towards jugum, JA generally smooth, or in some cases irregularly covered by converging longitudinal riblets. Articulamentum with apophyses short and wide, rounded, jugal sinus not very wide, almost straight to slightly concave, insertion lamina divided into short and rough denticles of irregular width, slit formula 7 / 1 / 8 – 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	discussion	Remarks. The fossil record of Stenosemus juliuspisai sp. nov. appears limited to the Mediterranean Pliocene (Italy and Spain) and Pleistocene (Italy). Dell’Angelo et al. (1999), in their study of the chitons from the Miocene (Tortonian) of Borelli (Italy), described some valves of the genus Stenosemus [as Ischnochiton (Simplischnochiton) exaratus] now attributed to S. praedolii sp. nov. (see below), illustrating also an intermediate valve from Altavilla Milicia (Dell’Angelo et al. 1999: pl. 3, fig. 6) which is attributable to S. juliuspisai sp. nov. The studied material could also be attributed to the genus Ischnochiton Gray, 1847, the generic attribution to Ischnochiton or Stenosemus is mainly based on the characters of the soft parts, e. g., the dorsal girdle covered with conical scales in Stenosemus, with imbricating, generally striated scales in Ischnochiton (Kaas & Van Belle 1990). The attribution to the genus Stenosemus is tentative, based on the great similarity of the valves here discussed with valves pertaining to Stenosemus [e. g., S. dolii (Van Belle & Dell’Angelo, 1998), S. vanbellei (Kaas, 1985), S. exaratus (G. O. Sars, 1878)]. The nodules present along the radial striae of the lateral areas on intermediate valves are highly variable in shape, size, and density; however, they are consistently distinct, elevated, and well separated from one another. We also provisionally include the intermediate valve from the Pliocene / Pleistocene of Estepona, Spain — figured in Dell’Angelo et al. (2004: pl. 7, figs 2, 5, 6) as Ischnochiton (Stenosemus) aff. vanbellei Kaas, 1985 — within the morphological range of Stenosemus juliuspisai sp. nov. Although the sculpture of the lateral areas appears different at first glance, we consider this variation to fall within the species’ variability and await additional material to confirm the precise taxonomic attribution. Some extant single intermediate valves of Stenosemus juliuspisai sp. nov. are present in the BD collection (currently under study, unpublished), in samples of debris collected by fishing boats, from Alghero (- 180 m, coral bottom) and Alboran Island (- 80 / 150 m). Some of these valves are illustrated for comparison (Figs 76 M – P). No significant differences were found between the fossil and extant valves described in this study. Comparisons. The intermediate valves of Stenosemus juliuspisai sp. nov. with only two radial striae of nodules present in the lateral areas show at first sight some similarities with S. vanbellei (a rare species randomly found in the Mediterranean Sea, not known as fossil), but differ mainly in the sculpture of the central area (11 – 16 longitudinal riblets on each side, normally all reaching anterior margin, vs. 7 – 9 longitudinal sulci in S. juliuspisai sp. nov., a little sinuous, the three or four innermost sulci not reaching the anterior margin).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF714EB00FADFCEF6945900C.taxon	distribution	Distribution. Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Rio Sant’Antonino, Zinola (Laghi 1977; Dell’Angelo et al. 2013, 2021 b; this study). Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Altavilla Milicia (Dell’Angelo et al. 2012), Valle Botto, Cava di Campore, Rio Merli (this study). Pleistocene: central Mediterranean, Italy: Capraia Island-Capo Corso 350 - 500 m (Dell’Angelo & Giusti, 1997), San Procopio (this study). Recent: Mediterranean Sea, Italy: single valves from Alghero and Alboran Island (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	description	Fig. 77	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	materials_examined	Type material. Holotype: MZB 32039, intermediate valve, width 5.7 mm (Figs 77 A – D). Paratype 1: MNHN. F. A 98475, intermediate valve, width 4.8 mm (Fig. 77 E) from Borelli (Italy), Miocene (Tortonian). Paratype 2: SMF 380820, intermediate valve, width 4.8 mm from Borelli (Italy), Miocene (Tortonian). Paratype 3: MSNG 62654, intermediate valve, width 5.6 mm (Figs 77 F – G) from Archi (Italy), lower Pleistocene. Paratype 4: MSNG 62655, intermediate valve, width 4.6 mm (Fig. 77 H) from Archi (Italy), lower Pleistocene. Type locality. Montegibbio (Emilia-Romagna, Italy). Type stage. Miocene (Tortonian).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	etymology	Etymology. The name recalls the morphological resemblance with the extant Mediterranean species Stenosemus dolii (Van Belle & Dell’Angelo, 1998).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	materials_examined	Material examined. Miocene (Tortonian). N. Italy: Borelli: 5 valves (BD 561, MZB 32041, MGPT PU 135043), Montegibbio: Holotype plus 1 valve (BD 562), Rio di Bocca d’Asino: 1 valve (BD 563). Lower Pleistocene. Italy: Calabria: Archi: 3 valves (BD 564). Maximum width of the intermediate valves: 8.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	diagnosis	Diagnosis. Intermediate valves broadly rectangular, width more than three times length, carinate, moderately elevated, apex inconspicuous. Tegmentum rough, sculptured with radial striae of small, roundish, elevated, clearly separated granules in LA, longitudinal riblets on each side in CA, more irregular and converging in JA. Articulamentum with apophyses short and wide, insertion lamina with a single slit.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	description	Description. Head and tail valves unknown. Intermediate valves broadly rectangular, width more than three times length (W / L = 3.07 – 3.18), carinate in anterior profile, moderately elevated (H / W = 0.37 – 0.50, Holotype 0.46), anterior margin consisting of three segments, two lateral ones straight or slightly concave, and one between apophyses straight or slightly convex, side margins almost straight or slightly rounded, posterior margin practically straight with mucro not very visible, lateral areas moderately elevated. Tegmentum rough. LA sculptured with 4 – 8 radial striae of small, roundish, elevated, clearly separated granules. CA sculptured with 15 – 20 or more longitudinal riblets on each side, normally all reaching anterior margin, more irregular and converging towards jugum, jugal area irregularly covered by converging longitudinal riblets. Articulamentum with apophyses short and wide, rounded, jugal sinus not very wide, slightly concave, insertion lamina with a single slit.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	discussion	Remarks. The fossil record of Stenosemus praedolii sp. nov. is limited to the Mediterranean upper Miocene and lower Pleistocene of Italy. Only intermediate valves are present in the material examined. The attribution to the genus Stenosemus is tentative, and the same considerations already expressed for Stenosemus juliuspisai sp. nov. apply (see above). Comparisons. The more similar species is Stenosemus juliuspisai sp. nov. (see above), from which Stenosemus praedolii sp. nov. differs mainly by the radial sculpture of the lateral areas of the intermediate valves, with 2 – 4 radial striae of large, quadrangular nodules in S. juliuspisai sp. nov., vs. 5 – 8 of radial striae of smaller, roundish and spaced granules in S. praedolii sp. nov.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4F4EB10FADFA116A9192BF.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N. Italy: Borelli, Montegibbio, Rio di Bocca d’Asino (Dell’Angelo et al. 2015 a). Lower Pleistocene: central Mediterranean, S. Italy: Archi (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4D4EB30FADFF0568C4947C.taxon	description	Fig. 78	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4D4EB30FADFF0568C4947C.taxon	materials_examined	Type material. Holotype MZB 50568, intermediate valve, width 5.4 mm, from Carrière Vives (France, Figs 78 C – E). Paratype MHNBx 2020.6.1, intermediate valve, from Lahitet (France). Type locality. Meilhan (France). Type stage. Miocene (Burdigalian). Material examined. Lower Miocene (Burdigalian): France: type material plus Carrière Vives: 7 valves (AC, Figs 78 F – H, BD 565, Figs 78 A – B). Maximum width of the valves: 4.5 / 5.4 / 4.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4D4EB30FADFF0568C4947C.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, slope almost straight. Intermediate valves broadly rectangular, width more than three times length, W / L = 3.00 – 3.71 (3.29 in holotype), semicarinate in anterior profile, moderately elevated (H / W = 0.25 – 0.41), anterior margin almost straight or slightly sinuose, side margins fairly straight, posterior margin almost straight at both sides of a not very prominent apex, lateral areas scarcely differentiated. Tail valve semicircular, anterior margin convex, mucro in anterior position, antemucronal slope slightly convex, postmucronal slope concave. Tegmentum rough. HV, LA and PMA sculptured with large and rhomboidal granules randomly arranged. CA and AMA sculptured with small and roundish granules randomly arranged. Articulamentum with large apophyses, slit rays numerous (more than 30 in head and tail valves, 4 – 5 in intermediate valves) and well evident in all valves, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4D4EB30FADFF0568C4947C.taxon	discussion	Remarks. The fossil record of Stenosemus radiatus Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 is limited to the lower Miocene of northeastern Atlantic and to the Middle Miocene of Paratethys. Stenosemus radiatus has two unusual characteristics, i. e., the intermediate valves being more than three times as wide as long, and the presence of a great number of slits in the insertion laminae. The slits are not always evident because of the worn condition of the valves, but the articulamentum displays numerous slit rays, and consequently an equal number of incisions on the insertion lamina should be expected. Also the generic attribution has been difficult, many extant species with a tegmentum entirely covered by granules arranged randomly are attributed to the genera Ischnochiton Gray, 1847, Stenosemus von Middendorff, 1847 or Lepidochitona Gray, 1821, mainly on the basis of the soft parts. Dell’Angelo et al. (2020 b) attributed the studied material to the genus Stenosemus, following Ferreira (1981) who considered the great number of slits in the insertion laminae as a character typical for this genus. Comparisons. Stenosemus radiatus differs from the other species of Stenosemus here described by the different sculpture (CA and AMA granulose in S. radiatus vs. longitudinal riblets in the other Stenosemus spp.), the shape of the intermediate valves and the higher number of slits (see Tab. 12).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4D4EB30FADFF0568C4947C.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Carrière Vives, Saint-Jean-de-Marsacq (Dell’Angelo et al. 2020 b). Middle Miocene: Central Paratethys (Langhian-Serravallian): Hungary: Borsodbóta (Schwabe & Dulai 2024).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	description	Fig. 79	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	materials_examined	Type material. Holotype: MSNG 62656, intermediate valve, width 7.7 mm (Figs 79 A – D). Paratype 1: MSNG 62657, intermediate valve, width 7.5 mm (Figs 79 E – H). Type locality. Pezzo (Calabria, Italy). Type stage. Upper Pleistocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	etymology	Etymology. The species is named after Antonietta Rosso (University of Catania) in recognition of her many paleontological and biological contributions on Mediterranean Cenozoic to Recent marine invertebrates.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	materials_examined	Material examined. Upper Pleistocene: Italy: Pezzo: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	diagnosis	Diagnosis. Intermediate valve broadly rectangular, carinate in anterior profile, elevated, apex small, LA delimited from CA by diagonal fold. Tegmentum smooth and glossy, with well marked concentric growth lines in CA, continuing across LA. Articulamentum well developed, with apophyses connected across shallow sinus by short jugal plate, insertion lamina with a single slit.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	description	Description. Head and tail valves unknown. Intermediate valve broadly rectangular (W / L = 2.25), carinate in anterior profile, elevated (H / W = 0.48), anterior margin almost straight or slightly convex, side margins almost straight, posterior margin weakly concave at both sides of small apex, lateral areas slightly raised, delimited from central area by diagonal fold. Tegmentum smooth and glossy to naked eye. CA with several well marked concentric growth lines, continuing, though less strongly, across LA. Articulamentum well developed, with apophyses rounded, connected across shallow sinus by a short, straight jugal plate, faintly notched at sides, insertion plates long with a slit, slit rays weakly indicated, teeth sharp, slightly rugose on outside, eaves narrow.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	discussion	Remarks. The fossil record of Stenosemus rossoae sp. nov. is limited to the upper Pleistocene of Italy. Two intermediate valves are present in the material studied, in good condition even if a little abraded. The generic attribution is difficult, the intermediate valves with a jugal plate separated from apophyses by small notches are typical of the genus Lepidozona Pilsbry, 1892, which, however, includes species with a tegmentum sculpture characterized by longitudinal riblets / radial rows of pustules (Sirenko 2021 b), not smooth or microgranulose. Other genera show the apophyses connected across the jugal portion, but all these have characteristics not agreeing with the material here studied, e. g., Callochiton Gray, 1847 (with a finely granulose sculpture, eaves spongy, extrapigmentary eyes present), Connexochiton Kaas, 1979 (of small size with a granulose sculpture), Stenosemus von Midendorff, 1847 (microgranulose). We provisionally attribute the material studied to Stenosemus, pending further material. The valve’s characters are distinctive and well defined, and for these reasons, and also considering the rare records of this genus as a fossil, we have described these two intermediate valves as a new species. Comparisons. Stenosemus rossoae sp. nov. is superficially similar to the living arctic-boreal species Stenosemus albus (Linnaeus, 1767), also reported from the Pleistocene of northeastern Atlantic (Antevs 1917, 1928; Feyling-Hanssen 1955), from which it differs by the smooth tegmentum (uniformly microgranulose in S. albus), LA delimited from CA by a diagonal fold (not present in S. albus), the greater dimensions of the valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4C4EB40FADFE016EF797B8.taxon	distribution	Distribution. Upper Pleistocene: central Mediterranean, S. Italy: Pezzo (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	description	Fig. 80	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	materials_examined	Type material. Holotype: MSNG 62658, tail valve, width 4 mm (Figs 80 A – D). Paratype 1: MSNG 62659, intermediate valve, width 3.2 mm (Figs 80 E – F). Paratype 2: MSNG 62660, head valve, width 4.3 mm (Figs 80 GH). Type locality. Borelli (Piedmont, Italy). Type stage. Miocene (Tortonian).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	etymology	Etymology. The name honors Michael J. Vendrasco (Pasadena City College, U. S. A.), for his prominent contribution to the study of Recent and fossil chitons.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	materials_examined	Material examined. Miocene (Tortonian): Italy: Borelli: type material. Maximum width of the valves: 4.3 / -- / 4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	diagnosis	Diagnosis. Head valve semicircular, tail valve semicircular, mucro not very prominent in anterior position, AMA short. Tegmentum granulose, sculptured with radial irregular striae of roundish granules in HV, PMA, and with longitudinal irregular granulose ribs in AMA. Articulamentum with apophyses short and wide, insertion lamina with 7 slits in holotype.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	description	Description. Intermediate valves unknown. Head valve semicircular, posterior margin widely V-shaped, with a rounded notch at apex. Tail valve semicircular, width more twice length (W / L = 2.13), anterior margin straight, more elevated and slightly concave between apophyses, mucro not very prominent in anterior position, AMA short, antemucronal and postmucronal slopes almost straight. Tegmentum granulose. HV and PMA sculptured with radial irregular striae of roundish granules (ca. 40 in PMA), obsolete towards apex or mucro, with evidence of irregularly arranged granules. AMA sculptured with longitudinal irregular granulose ribs, also in JA. Articulamentum with apophyses short and wide, insertion lamina divided into short and rough denticles of irregular width, 7 slits in holotype.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	discussion	Remarks. The fossil record of Stenosemus vendrascoi sp. nov. is limited to the Mediterranean upper Miocene of Italy. Only one head and two tail valves are present in the material examined. Also for this species the attribution to the genus Stenosemus is tentative, and the same considerations already expressed for Stenosemus juliuspisai sp. nov. apply (see above). The shape of the tail valve designed as paratype shows some difference in the antemucronal (slightly convex) and postmucronal (slightly concave just under mucro), but the other characters (tegmentum sculpture, W / L, AMA short) agree, and we consider these variations to be part of intraspecific variability. Comparisons. See Tab. 12 for a comparison with the Stenosemus spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N. Italy: Borelli (this study). Genus Connexochiton Kaas, 1979 = Bathychiton Dell’Angelo & Palazzi, 1988	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	type_taxon	Type species. Connexochiton platynomenus Kaas, 1979, by original designation.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	distribution	Distribution. Connexochiton is known from Pleistocene to Recent, with six living species known from the Atlantic, Indian and Pacific Oceans, and the Mediterranean Sea (Sirenko 2021 a). The fossil record includes the lower Miocene of France (Dell’Angelo et al. 2018 a) and the Pliocene-Pleistocene of Italy (Dell’Angelo & Palazzi 1988, 1994; Dell’Angelo et al. 2013).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4B4EB50FADFD4568229016.taxon	discussion	Remarks. The genus Connexochiton Kaas, 1979 has been recently revised by Sirenko (2021 a). The main morphological characters of the Connexochiton spp. considered in the present study are reported in Tab. 11, together with the species of the genus Stenoplax.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	description	Fig. 81	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	materials_examined	Type material. Holotype MNHN, specimen partly curled, length 2.3 mm, width 1.6 mm. Paratype RMNH K 4814. Holotype of Bathychiton biondii Dell’Angelo & Palazzi, 1988 at MSNL, disarticulate specimen, ca. 5 x 3 mm (Figs 81 G – H). Type locality. Bay of Biscay, 800 – 1050 m. Material examined. Pleistocene. Italy: Calabria: Archi: 2 valves (BD 566, Figs 81 A – D), Bovetto: 1 valve (BD 567, broken), Musalà: 2 valves (BD 568), Vallone Catrica, 1 valve (BD 569, Figs 81 E – F); Sicily: Gravitelli: 3 valves (BD 570). Recent. Italy: type material of Bathychiton biondii. Maximum width of the valves: 2 / 2.8 / 1.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	description	Description. Head valve semicircular, highly elevated, posterior margin somewhat sinuate. Intermediate valves broadly rectangular, ii – iii highly elevated (H / W = 0.67 – 0.75), carinate in anterior profile, rather long (W / L = 1.50 – 1.90), anterior margin strongly convex, valves iv – vii less elevated (H / W = 0.43), semicarinate in anterior profile, much shorter (W / L = 2.40 – 3.00 or more), anterior margin straight or slightly concave, all valves with lateral areas distinctly raised. Tail valve semicircular, nearly twice as wide as long (W / L = 1.88 – 2.03), anterior margin almost straight, mucro subcentral, not prominent, postmucronal slope slightly concave. Tegmentum rough, sculptured with relatively large, roundish, flat-topped granules (diameter 75 µ), neatly arranged in quincunx, as a result they are also directed in radial rows on LA; each granule with 10 – 12 aesthetes of equal size, irregularly distributed. Some intermediate and tail valves show a narrow “ pre-central area ” adjacent to anterior margin coarse without granules. Articulamentum with apophyses somewhat triangular in shape, rectangular in tail valve, jugal sinus flat and shallow as a result of a considerable laminate extension of articulamentum, connecting apophyses, jugal plates very slightly notched, notches corresponding with faint grooves, not separated from apophyses by prominent slits, insertion plates very short, slit formula 11 / 1 / 9 – 10, teeth decidedly propped, especially in tail valve, eaves narrow, solid.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	discussion	Remarks. Connexochiton platynomenus Kaas, 1979 has been described on a small juv. (3 mm long) incomplete (valves vii-viii lacking) specimen from the Bay of Biscay. Dell’Angelo & Palazzi (1998) described Bathychiton biondii on a larger specimen (5 mm long), on the basis of some characters not present (or not described) in C. platynomenus, among which the presence of a “ pre-central area ” without granules in the intermediate and tail valves and the presence of a row of spherules on the top of dorsal scales. The presence of the pre-central area seems to be quite variable in the scarce material examined, from clearly evident (Figs 76 E – G) to absent (Fig. 76 C), while subsequently Kaas & Van Belle (1990), after a careful reexamination of the girdle of the holotype of C. platynomenus, found a few girdle scales with a marginal row of spherules, and speculated that, as the holotype of C. platynomenus is 50 % smaller that the holotype of B. biondii, it is possible that the spherules appear at a later stage of growth. Fossil records of these species are scarce, therefore we prefer to consider B. biondii a synonym of C. platynomenus, awaiting further material (both fossil and living) to study. Kaas (1979) already highlighted in the original description the difference between the first two intermediate valves (ii – iii) and the subsequent ones (iv – vii), as regards shape and elevation. These differences, clearly visible by analyzing the plates of a whole specimen, are also easily recognizable in the single intermediate valves found as fossils (e. g., Figs 76 C – D for valves ii – iii and Figs 76 E – F for valves iv – vii). Comparisons. See Tab. 11 for a comparison with the Connexochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	distribution	Distribution. Pleistocene: central Mediterranean, S. Italy: Archi, Bovetto, Gravitelli, Musalà, Vallone Catrica (Dell’Angelo & Palazzi 1988, 1994; this study). Recent: Atlantic Ocean: Atlantic side of Strait of Gibraltar: Bay of Biscay; Canary Islands: SW of Hierro Island (Kaas 1979; Kaas & Van Belle 1988); Mediterranean Sea: S. Lucia Bank, Tuscan Archipelago (Dell’Angelo & Palazzi 1988). “ Connexochiton ” roccai Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013 Fig. 82	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	materials_examined	Type material. Holotype MZB 49980, tail valve, width 1.7 mm, Figs 82 A – D. Type locality. Borzoli (Liguria, ltaly). Type stage. Lower Pliocene (Zanclean). Material examined. Lower Pliocene: Italy: Borzoli: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	description	Description. Head and intermediate valves unknown. Tail valve small, semicircular, rather flat, mucro situated anteriorly, not prominent, antemucronal and postmucronal slopes almost straight. Tegmentum rather coarsely granulose. PMA sculptured with roundish granules united with each other (diameter 48 – 53 µm), forming fine network. AMA sculptured with granules arranged in segments of various size and shape, slightly overlapping each other to form very irregular longitudinal rugosities, obliquely oriented with respect to anterior-posterior axis. Apophyses rounded, wide, connected across jugal portion, insertion plate thick, eaves solid, teeth rough, irregular, 10 slits visible, total number estimated at 12.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	discussion	Remarks. “ Connexochiton ” roccai Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013 is a very rare taxon, known only from a single tail valve found in the Lower Pliocene of Borzoli. The valve’s characters are very distinctive and well defined, to justify the description of a new species. Until more material is discovered, variability remains unknown. The generic attribution is difficult, and the emended diagnosis of the genus Connexochiton by Sirenko (2021 a) does not fully meet the characteristics of the valve of “ Connexochiton ” roccai, as regards the sculpture of the tegmentum. This species is provisionally referred to Connexochiton based on articulamentum characters, though the tegmentum sculpture deviates from the genus diagnosis. According to Sirenko (2021 a), the genus Connexochiton superficially resembles five genera: Subterenochiton Iredale & Hull, 1924, Thermochiton Saito & Okutani, 1990, Callistochiton Carpenter MS, Dall, 1879, Stenosemus Middendorff, 1847 and Lepidozona Pilsbry, 1892. We provisionally attribute the material studied to Connexochiton, pending the avalability further material. Comparisons. See Tab. 11 for a comparison with the Connexochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	distribution	Distribution. Lower Pliocene: central Mediterranean, Italy: Borzoli (Dell’Angelo et al. 2013, 2021 b). “ Connexochiton ” vivesi Dell’Angelo, Lesport, Cluzaud & Sosso, 2018 Fig. 83	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	materials_examined	Type material. Holotype: MHNBx 2017.6.1, tail valve, width 4 mm (Figs 83 A – D). Type locality. Meilhan, Carriere Vives (France). Type stage. Lower Miocene (Burdigalian). Material examined. Lower Miocene: France: Meilhan: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	description	Description. Head and intermediate valves unknown. Tail valve small, semicircular, length about half width, rather flat, anterior margin slightly convex, mucro situated anteriorly, not prominent, antemucronal slope almost straight, postmucronal slope slightly concave just underneath mucro and quite straight posterior from there. Tegmentum rather coarsely granulose. PMA sculptured with granules arranged in segments of various size and orientation, slightly overlapping each other forming rugosities. AMA sculptured with roundish granules fused forming fine network. Apophyses rounded, wide, connected across jugal portion, insertion plate thick, eaves solid, central depression with numerous transverse slits in jugal tract, teeth eroded and scarcely visible, but 28 well marked slits rays are present.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	discussion	Remarks. “ Connexochiton ” vivesi Dell’Angelo, Lesport, Cluzaud & Sosso, 2018 is known only for a single tail valve from the Lower Miocene of Meilhan. The valve is partly eroded, also the apophyses are not complete, but the connection by a lamina is still visible. Also the posterior margin is eroded, the incisions are barely outlined and hardly visible. Nevertheless, the valve’s characters are very distinctive and well defined, and justify the description of a new species. Until more material is discovered, variability remains unknown. The generic attribution is difficult, since the emended diagnosis of the genus Connexochiton by Sirenko (2021 a) does not fully meet the characteristics of the valve of “ Connexochiton ” vivesi, as regards the sculpture of the tegmentum. This species is provisionally referred to Connexochiton based on articulamentum characters, though the tegmentum sculpture deviates from the genus diagnosis. Comparisons. “ Connexochiton roccai ” Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013 has a granulose sculpture with some resemblance to C. vivesi, but the pattern is opposite, with the network in postmucronal area and the irregular concentric wrinkles in antemucronal one (see Tab. 11).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF4A4EB80FADFAEE6BBC93C4.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (early Burdigalian): Aquitaine Basin, France: Meilhan (Dell’Angelo et al. 2018 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF474EB90FADF8D86B5A9743.taxon	type_taxon	Type species. Callistochiton palmulatus Carpenter MS, Dall, 1879, by monotypy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF474EB90FADF8D86B5A9743.taxon	distribution	Distribution. Callistochiton is known from the Oligocene to the Recent. The genus Callistochiton is widespread, occurring in cool to warm waters worldwide (Kaas & Van Belle 1994). The sole NE Atlantic and Mediterranean species is C. (Allerychiton) pachylasmae (Monterosato, 1879). The fossil record for Callistochiton is largely incomplete (Puchalski et al. 2008), and the few records do not allow us to suggest a diversification scenario for the group. The fossil record extends back to the Oligocene deposits of Germany (Janssen 1978; Dell’Angelo et al. 2011) and New Zealand (Lee et al. 2014), the Miocene of Japan (Itoigawa et al. 1981) and Tanzania, E. Africa (Davis 1954), the Miocene-Pliocene of California, U. S. A. (Vendrasco et al. 2012, 2022 and references therein), the Pliocene-Pleistocene of S. Italy (Dell’Angelo et al. 1998 b), and the Pleistocene (interglacial deposits) of California, U. S. A. (Muhs et al. 2002).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF474EB90FADF8D86B5A9743.taxon	discussion	Remarks. The main morphological characters of the Callistochiton spp. considered in the present study are reported in Tab. 13333. observations).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF464EBA0FADF8CD6B5F900A.taxon	description	Fig. 84	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF464EBA0FADF8CD6B5F900A.taxon	materials_examined	Type material. Holotype MGPT PU 108786, intermediate valve, width 5.7 mm (Figs 84 A – D). Type locality. Borelli (Piedmont, Italy). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian). Italy: Borelli: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF464EBA0FADF8CD6B5F900A.taxon	description	Description. Head and tail valves unknown. Intermediate valve solid, broadly rectangular (W / L = 1.98 estimated), carinate in anterior profile, elevated (H / W = 0.61), anterior and posterior margins straight, side margins rounded, slightly bilobated, apices inconspicuous, lateral areas raised, distinctly defined. LA sculptured with two nodulose ribs of different width, rib near the diagonal ridge wider, crossed by numerous concentric grooves, and separated by narrow, deep sulcus. CA sculptured with 11 longitudinal, elevated, granulose riblets, becoming larger towards the lateral margins, JA large, smooth, only marked by some growth lines. Articulamentum with apophyses wide, insertion plates well developed, apical area prominent, teeth not pectinated, a single short slit for each side, slit rays hardly discernible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF464EBA0FADF8CD6B5F900A.taxon	discussion	Remarks. Callistochiton borellianus Dell’Angelo, Giuntelli, Sosso & Zunino, 2015 is known only for a unique intermediate valve from the upper Miocene of N. Italy. The valve is partly eroded, but the diagnostic characters are very distinctive and well defined. The unique intermediate valve has been assigned to the genus Callistochiton on the basis of strong similarities to species included in this genus (Dell’Angelo et al. 2015 a), and represents the first record of this genus for the Italian Miocene. Comparisons. See Tab. 13 for a comparison with the Callistochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF464EBA0FADF8CD6B5F900A.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Borelli (Dell’Angelo et al. 2015 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF454EBC0FADFA1A680A95C4.taxon	description	Fig. 85	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF454EBC0FADFA1A680A95C4.taxon	description	121, pls 39, 39 [bis], fig. 54; Dell’Angelo et al. 2015 a, p. 238; Dell’Angelo et al. 2018 c, p. 216, figs 1 – 3. Callistochiton (Allerychiton) pachyplasmae [sic]; Koukouras & Karachle 2005, p. 30. Callistochiton sp. Dell’Angelo et al. 1998 b, p. 139, fig. 2; Dell’Angelo & Smriglio 2009, p. 124; Dell’Angelo et al. 2015 a, p. 238 (fide Dell’Angelo et al. 2018 c).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF454EBC0FADFA1A680A95C4.taxon	materials_examined	Type material. Holotype: MCZR – M – 12683 / H, a specimen (3.4 x 1.3 mm) collected by G. Seguenza on Pachylasma giganteum (Appolloni et al. 2018). Type locality. Strait of Messina (Italy). Material examined. Pleistocene: Italy: Calabria: Santa Maria di Catanzaro; 1 head valve, width 1.3 mm (BD 571, Figs 85 A – B); Gallina: 1 head valve, width 1.5 mm (AV, Figs 85 C – D), Pecoraro: 1 intermediate valve, width 1.6 mm (BD 572, Figs 85 G – H). Recent: Italy: Pantelleria (SB), Fig. 80 F; Greece: Corfu (SB), Fig. 85 E.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF454EBC0FADFA1A680A95C4.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, with a rounded apical notch. Intermediate valves broadly rectangular, elevated, semicarinate in anterior profile, posterior margin nearly straight and apex slightly prominent, lateral areas raised. Tail valve semicircular, mucro slightly posterior, not prominent, postmucronal slope slightly concave. HV and LA sculptured with large wrinkled, elevated radial ribs, 6 – 8 in HV starting from edge of apparently smooth apical area, interstices coarsely granulated, one in LA, starting from apex, covered by scaly tubercles. PMA with one wrinkled radial rib on either side a little behind antemucronal-postmucronal boundary, sculptured with large tubercles irregularly disposed, appearing arranged along concentric lines. CA and AMA sculptured with large, irregularly disposed, close-set tubercles, surface between tubercles strongly coarse, tubercles irregularly roundish, larger diameter ca. 44 – 56 µm in CA, with up to three aesthetes in lower part and another one or two slightly larger in more central position. Tail valve with antemucronal area sculptured with large tubercles as in central area of intermediate valves, Articulamentum perlaceous, apophyses small and triangular, intermediate valves with two deep lateral dimples near apical area, with a marked radial grove, insertion plate divided by one deep cut on either side in two smooth teeth, slit formula 6 – 8 / 1 /?.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF454EBC0FADFA1A680A95C4.taxon	discussion	Remarks. The name was made available by a description included in the paper of Monterosato on Polyplacophora (1879), but the manuscript name by Giuseppe Seguenza already appeared as a nomen nudum in Monterosato (1878 a: 77) and Tiberi (1879: 150). Callistochiton pachylasmae (Monterosato, 1879) is the type species of the subgenus Allerychiton Dell’Angelo & Oliverio, 1997 (by original designation), proposed to highlight the unusual sculpture of this species, whose tail valve is different from that of the head valve, and with a single radial rib present on the lateral areas of the intermediate valves, vs. 2 or more ribs present in the other Callistochiton species (Kaas & Van Belle 1994). Dell’Angelo et al. (2018 c) reported the second documented record of C. pachylasmae from the Strait of Messina, of this rarely recorded species. These authors contributed to the knowledge of habitat and distribution of this enigmatic species. The fossil record of Callistochiton pachylasmae is limited to the Pleistocene (Calabrian) of southern Italy, with two head valves from Gallina (Reggio Calabria) and Santa Maria di Catanzaro, determined by Dell’Angelo et al. (1998 b) as Callistochiton sp. and C. pachylasmae, respectively, and an intermediate valve from Pecoraro. Comparisons. Callistochiton pachylasmae has CA sculptured with large, irregularly disposed, close-set tubercles, and this is the main difference towards the other two Callistochiton species discussed in this work, both showing a sculpture of granulose longitudinal ribs (see Tab. 13).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF454EBC0FADFA1A680A95C4.taxon	distribution	Distribution. Pleistocene: central Mediterranean, southern Italy: Gallina, Pecoraro, Santa Maria di Catanzaro (Dell’Angelo et al. 1998 b; this paper). Recent: Atlantic Ocean: Morocco (Kaas, 1991); Mediterranean Sea: Italy, Croatia, Greece (Dell’Angelo et al. 2018 c).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF434EBD0FADFED86BAC9490.taxon	description	Fig. 86	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF434EBD0FADFED86BAC9490.taxon	materials_examined	Type material. Lectotype SMF 14.28 a (Fig. 86 H) designed by Janssen 1978, tail valve figured by Boettger (1869), pl. 1 b, figs 13 a – b (Fig. 86 E). Paralectotypes SMF 14.28 b – c (Figs 86 F – G), valves figured by Boettger (1869), pl. 1 b, figs 13 c – e (Figs 86 C – D) and 13 f – g (Figs 86 A – B), respectively. Type locality. Gienberg (Germany). Type stage. Oligocene (Rupelian). Material examined. No actual material available, only descriptions and illustrations from the literature.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF434EBD0FADFED86BAC9490.taxon	description	Description. Head valve semicircular, strongly arched, posterior margin widely V-shaped. Intermediate valve broadly rectangular (W / L = 2.14), moderately elevated, semicarinate in anterior profile, anterior margin slightly sinuose at both sides of jugal area slightly extended forward, side margins slightly rounded, posterior margin almost straight at both sides of well evident apex, lateral areas not raised. Tail valve semicircular / elliptical (W / L = 1.68), mucro small, subcentral. HV, LA and PMA sculptured with radial granulose ribs (HV 11 – 14, with some finer secondary ribs, LA 3 – 4, PMA 11 – 18). PA and AMA sculptured with granulose longitudinal riblets (CA 7 – 12 on half valve, usually 9), crossed by few concentric ridges, JA smooth. Articulamentum with apophyses very large, broad, protruding far, connected in middle, leaving only a narrow sinus free, apical area in intermediate valves triangular and small, insertion plates short, slit formula 25 / 3 – 4 / 15, teeth strongly furrowed on upper side and thickened at edges.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF434EBD0FADFED86BAC9490.taxon	discussion	Remarks. The fossil record of Callistochiton zitteli (de Rochebrune, 1882) is limited to the Oligocene (Rupelian) of Germany. Chiton fimbriatus Boettger, 1869 could not be mantained as it was already known Chiton fimbriatus Sowerby, 1840, a species from Peru now considered “ taxon inquirendum ” by WoRMS. Comparisons. Callistochiton zitteli is characterized by a number of slits far greater than the other Callistochiton discussed in this study (see Tab. 13).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF434EBD0FADFED86BAC9490.taxon	distribution	Distribution. Lower Oligocene: North Europe: Mainz Basin, Germany: Gienberg, Heimberg, Zeilstück (Janssen 1978; Hocht 1986).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF424EBD0FADFDD46AB79068.taxon	type_taxon	Type species. Chiton affinis Issel, 1869, by subsequent designation (International Committee of Zoological Nomenclature 1971, Opinion 951, as proposed by Beu et al. 1969).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF424EBD0FADFDD46AB79068.taxon	distribution	Distribution. Rhyssoplax is reported from the Oligocene up to the Recent, with a present distribution in the Indo-West Pacific, and eastern Atlantic Ocean, Mediterranean included. The fossil record encompasses the Oligocene of New Zealand (Lee et al. 2014; Wu & Lee 2024), Neogene of Europe (Dell’Angelo et al. 2004, 2015 a; Garilli et al. 2005; Studencka & Dulai 2010), Africa (Algeria: De Lamothe 1911; Tunisia: Castany et al. 1956), New Zealand (Beu & Maxwell 1990; Sutherland et al. 1995) and Japan (Itoigawa et al. 1976), Pleistocene of the Red Sea (Dell’Angelo et al. 2020 a) and Madeira (Gerber et al. 1989), Holocene of Japan (Kuroda et al. 1980).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF424EBD0FADFDD46AB79068.taxon	discussion	Remarks. The genus Chiton was subdivided into numerous subgenera by Kaas et al. (2006), some of which (e. g., Rhyssoplax Thiele, 1893) later erected to genus by Sirenko (2006), based mainly upon soft part morphology. We consider here ten species attributed to Rhyssoplax, whose characteristics are summarized in tables 14 and 15, conveniently arranged in two groups based upon differences in tegmentum sculpture of HV, LA and PA. The first group includes four species, with the presence of more or less pronounced radial ribs [R. miocenica (Michelotti, 1847), R. olivacea (Spengler, 1797), R. saeniensis (Laghi, 1984) and R. sulcomarginata (( Dell’Angelo, Giuntelli, Sosso & Zunino, 2016)] (Tab. 14). The second group includes 6 species [R. assurrecta Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018, R. capecchii (Chirli, 2004), R. corallina (Risso, 1826), R. etrusca (Dell’Angelo & Forli, 1995), R. garillii sp. nov. and R. phaseolina (Monterosato, 1879)], whose areas are instead smooth (Tab. 15).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF404EA10FADFA50698894C8.taxon	description	Fig. 87	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF404EA10FADFA50698894C8.taxon	materials_examined	Type material. Holotype MNHN. F. A 67124, intermediate valve, width 6.5 mm (Figs 87 E – H). Paratypes: MNHN. F. A 67125 – A 67128 (4 valves) (Figs 87 I – K); NHMW 2017 / 0108 / 0033 – 0036 (4 valves) (Figs 87 A – B); RGM. 1008368 – 1008371 (4 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Lower Miocene: France (Aquitanian): Maïnot: 1 valve (PR). France (Burdigalian): Carrière Vives: 1 valve (BD 639). Upper Miocene: France (Tortonian): Saint-Clément-de-la-Place: type material plus 230 valves (MNHN. F. A 67129 – A 67130, NHMW 2017 / 0108 / 0037, RGM. 1008355, RGM. 1008372, BD 151); Sceaux d’Anjou, La Presselière: 1 valve (RGM. 1008450). France (Messinian?): Moulin-Pochas: 1 valve (PR). Lower Pliocene: Italy: Borzoli: 37 valves (BD 640, MZB 45747); Bussana: 13 valves (BD 641); Caranchi: 5 valves (BD 642, MP); Rio S. Antonino: 4 valves (BD 643, MP). Pliocene: Italy: Tuscany: Pietrafitta Melograni: 10 valves (BD 644, Figs 87 C – D, 87 L), Poggio alla Fame: 12 valves (BD 645, MP). Maximum width of the valves: 5.7 / 7.5 / 4.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF404EA10FADFA50698894C8.taxon	description	Description. Head valve semicircular, posterior margin markedly V-shaped, front slope straight. Intermediate valves broadly rectangular (W / L = 1.90 – 2.35), carinate in anterior profile, elevated (H / W = 0.55 – 0.71), anterior margin slightly convex, side margins truncated, posterior margin straight, interrupted by small apex, lateral areas raised, clearly defined. Tail valve more than semicircular (W / L = 1.38 – 1.63), anterior margin convex, mucro subcentral, antemucronal slope slightly convex, postmucronal slope almost straight or slightly concave, just behind mucro. HV, LA and PMA smooth, concentric growth lines weakly developed, more distinct on head and tail valves. CA and AMA sculptured with up to 10 – 11 irregular, longitudinal ribs on both sides of broadly triangular, smooth jugum, 2 – 4 innermost ribs not reaching anterior margin. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with wide, rounded apophyses, teeth irregular, slit formula 8 / 1 / 10 – 12, slit rays conspicuous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF404EA10FADFA50698894C8.taxon	discussion	Remarks. This species has been attributed to the genus Rhyssoplax based on the morphological affinities with the morphological closer R. corallina (Risso, 1826). Valves reported from the Pliocene of Liguria as Chiton sp. A (Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013) have been later ascribed to R. assurrecta (Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018). Comparisons. Rhyssoplax assurrecta Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is very similar to R. corallina (Risso, 1826), from which it differs mainly by the more elevated intermediate valves, H / W = 0.55 – 0.71 vs. 0.30 – 0.47 in R. corallina. The elevated intermediate valves are also a constant character in juvenile specimens (Dell’Angelo et al. 2018 b, compare figs 19 K – L and 18 F). The head valves of Rhyssoplax assurrecta and R. corallina are similar, and can only be separated based on the higher elevation of the valves of R. assurrecta (Dell’Angelo et al. 2018 b, compare figs 19 F and 18 B).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF404EA10FADFA50698894C8.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic: Aquitaine Basin, France: Maïnot (Aquitanian), Carrière Vives (Burdigalian) (Dell’Angelo et al. 2018 a). Upper Miocene: northeastern Atlantic: Ligerian Basin, France (Tortonian): Saint-Clément-de-la-Place, Sceaux d’Anjou (Dell’Angelo et al. 2018 b); Ligerian Basin, France (Messinian?): Moulin Pochas (Dell’Angelo et al. 2018 a). Pliocene: central Mediterranean, Italy: Liguria (Dell’Angelo et al. 2013); Tuscany: Pietrafitta, Poggio alla Fame (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5E4EA20FADFDD46EDA91CB.taxon	description	Fig. 88	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5E4EA20FADFDD46EDA91CB.taxon	materials_examined	Type material. Holotype MZB 42947. Type locality. Poggio alla Fame, Siena (Tuscany, Italy). Type stage. Pliocene, “ sabbie argillose grigie ” (gray clayely sands). Material examined. Pliocene: Italy: Tuscany: Bibbiano: 12 valves (BD 646), Cetona: 2 valves (BD 647), Pietrafitta: 60 valves (BD 648), Pietrafitta Melograni: 46 valves (BD 649, Fig. 88 D), Poggio alla Fame: 20 valves (BD 650, Figs 88 A – C, 88 E – L). Maximum width of the valves: 2.8 – 3.3 – 2.6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5E4EA20FADFDD46EDA91CB.taxon	description	Description. Head valve semicircular, front slope slightly convex, posterior margin widely V-shaped. Intermediate valves broadly rectangular, elongated (W / L = 2.27 – 3.30), moderately elevated (H / W = 0.29 – 0.37), carinate in anterior profile, anterior margin straight, side margins almost straight, posterior margin straight, minute apex, lateral areas not raised and not separated from CA. Tail valve semicircular (W / L = 1.71 – 1.83), anterior margin straight, mucro in anterior position, antemucronal slope straight, postmucronal slope slightly concave. Tegmentum smooth, crossed by few growth lines just highlighted (not in LA). Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses tending to trapezoidal, jugal sinus large, slit formula 12 – 15 / 1 / 15 – 16 (a second slit ray lies close to posterior margin on intermediate valves), teeth short, irregular, finely pectinated, slits narrow, slit rays very finely indicated.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5E4EA20FADFDD46EDA91CB.taxon	discussion	Remarks. The fossil record of Rhyssoplax capecchii (Chirli, 2004) is thus far limited to the Pliocene of Tuscany (Italy). Chirli (2004) described the small valves of this species (up to 3.6 mm of width) and highlighted the similarity to Rhyssoplax saeniensis (Laghi, 1984); in his opinion, R. capecchii differs from its smaller size, and other characteristics (Chirli 2004), which could well be included in the known variability of shape and sculpture existing in growth series of Polyplacophora [see, for example, Dell’Angelo et al. 2012, for Stenosemus dolii (Van Belle & Dell’Angelo, 1998) from Altavilla; Dell’Angelo et al. 2022, for Ischnochiton zbyi Dell’Angelo & Silva, 2003 from Vale de Freixo]. Laghi (1984) already highlighted such valve’s variability, in the original description of R. saeniensis a growth series of head, intermediate and tail valves (Laghi 1977: pl. 1, figs 1 – 5, 7 – 13, 15 – 20, respectively). The main differences between Rhyssoplax capecchii and R. saeniensis stay, therefore, in the considerably smaller size (maximum valve’s width 3.3 mm for R. capecchii) and the smooth tegmentum of R. capecchii, whereas R. saeniensis consistently display little-pronounced flat radial / longitudinal ribs. Moreover, the two taxa apparently inhabited different habitats: R. saeniensis is more often associated with shallow water sandy-gravels (Forli et al. 2003), whilst R. capecchii lived in sandy-clayey bottoms (Chirli & Bogi 2002; Bogi & Chirli 2004). Despite the great quantity of valves of R. saeniensis examined from Serre di Rapolano (> 1500), the smallest valve found has a width of 5.1 mm. Further, the two species do not co-occur in the sites where they are most frequently found (Serre di Rapolano for R. saeniensis, Poggio alla Fame and Pietrafitta Melograni for R. capecchii). We prefer at this stage to keep, therefore, the two taxa distinct. Comparisons. Rhyssoplax capecchii differs from the other Rhyssoplax spp. mainly by the tegmentum entirely smooth.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5E4EA20FADFDD46EDA91CB.taxon	distribution	Distribution. Pliocene: central Mediterranean, Italy: Tuscany: Bibbiano, Cetona, Pietrafitta, Pietrafitta Melograni, Poggio alla Fame (Laghi 1984; Dell’Angelo et al. 2001 a; Chirli 2004; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5D4EA60FADFAD56A9495E8.taxon	description	Fig. 89	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5D4EA60FADFAD56A9495E8.taxon	materials_examined	Type material. Presumably lost, not present in the Risso collection hosted at MNHN (fide Arnaud 1977). Type locality. Nice (France). Material examined. Lower Miocene: France (Aquitanian): Maïnot: 3 valves (JFL); France (Burdigalian): Carrière Vives: 5 valves (BD 651), Coupe du fossé près de La Solitude: 2 valves (JFL, Figs 89 B – D), Lahitet: 13 valves (AC, JFL, PR); Italy (Burdigalian): Valle Ceppi: 17 valves (BD 652, MZB 32071 – 32073, Figs 89 J – L, PG), Sciolze: 10 valves (PG). Middle Miocene: Italy (Langhian): Albugnano: 2 valves (MZB 32074, PG); Central Paratethys (Langhian-Serravallian): Romania: Bujtur: 1 valve (NHMW 1863 / 0015 / 0116, as Chiton bohemicus, Figs 89 H – I), Costej: 3 valves (BD 653), Lăpugiu de Sus: 3 valves (BD 654), Hungary: Bánd: 34 valves (BD 655), Letkés: 18 valves (BD 656), Sámsonháza: 1 valve (BD 657), Várpalota: 1 valve (BD 658); Eastern Paratethys: Ukraine: Varovtsi: 2 valves (BD 659), Zalistsi: 1 valve (BD 660); France (Serravallian): Orthez (Le Paren): 2 valves (BD 661), Sallespisse (Carré): 1 valve (PR). Upper Miocene: Italy (Tortonian): Borelli: 1 valve (BD 662), Montegibbio: 8 valves (BD 663, MZB 32076 – 32077, Figs 89 E – F), Rio di Bocca d’Asino: 112 valves (BD 664, Fig. 89 A, MZB 32075, Figs 89 M – N, PG), Vargo: 2 valves (PG), Vigoleno: 11 valves (BD 665), Villa Monti: 40 valves (BD 666, PG); France (Tortonian): Beugnon: 1 valve (RGM. 1310176), Renauleau: 8 valves (BD 149, MNHN. F. A 67123), Saint-Clément-de-la-Place: 4009 valves (BD 148, Figs 89 O – P, MNHN. F. A 67117 – A 67122, NHMW 2017 / 0108 / 0032, RGM. 1008354, RGM. 1008367, RGM. 1008435), Sceaux d’Anjou: 17 valves (BD 150, RGM. 1008444, RGM. 1008449); France (Messinian?): Moulin-Pochas: 5 valves (PR). Lower Pliocene: Italy: Borzoli: 2788 valves (BD 667), Bussana: 795 valves (BD 668, PG, MZB 45733, MZB 45735 – 45736), Caranchi: 34 valves (BD 669, MP), Garlenda: 364 valves (MP), Genova Sestri: 69 valves (BD 670), Rio Sant’ Antonino: 2620 valves (BD 671, MP, PG, MZB 45730 – 45732, MZB 45734, MZB 45737 – 45742), Rio Torsero: 29 valves (BD 672, MP, PG), Salea: 2 valves (BD 673), Zinola: 87 valves (BD 674). Pliocene: Spain: Estepona: 250 valves (BD 675). Portugal: Vale de Freixo: 664 valves (BD 243, GeoFCUL VFX. 03.342 – 343, MNHN. F. A 81988 - A 81989, RGM. 1364014 - 1364015). Italy: Piedmont: Moncalvo: 11 valves (BD 676), San Damiano: 15 valves (BD 677), Valle Botto: 1 valve (BD 678); Emilia-Romagna: Castell’Arquato: 3 valves (BD 679), Cava di Campore: 768 valves (BD 680), Ceparano: 2 valves (BD 681), Gagliardella: 16 valves (BD 682), Marzeno: 1 valve (BD 683), Monte Castellaccio: 3 valves (BD 684), Pietra Mora: 1 valve (BD 685); Tuscany: Bibbiano: 30 valves (BD 686), Castell’Anselmo: 4 valves (BD 687), Castelnuovo Berardenga: 1 valve (BD 688), Castiglioncelio del Trinoro: 2 valves (BD 689), Cetona “ Palazzo Tosoni ”: 2 valves (BD 690), Cisternino: 2 valves (BD 691), Colle Val d’Elsa: 25 valves (BD 692), Montenero: 16 valves (BD 693), Orciano Pisano: 185 valves (BD 694), Pietrafitta: 19 valves (BD 695), Poggio alla Fame: 2 valves (BD 696, Fig. 89 G), Pontedera: 3 valves (BD 697), Serre di Rapolano: 27 valves (BD 698), Villa Banfi: 1 valve (BD 699); Latium: Magliano Sabina: 3 valves (BD 700); Sicily: Trappeto: 19 valves (BD 701), Sciacca: 18 valves (BD 702). Upper Pliocene to upper Pleistocene: France: Bosq d’Aubigny: 5 valves (RGM. 1310186). Pleistocene: Italy: Tuscany: Caletta: 4 valves (BD 703), Capraia Island-Capo Corso: 35 valves (BD 704), Cisternino: 5 valves (BD 705), Riparbella: 3 valves (BD 706); Puglia: Cutrofiano: 1 valve (BD 707), Gallipoli: 15 valves (BD 708); Calabria: Archi S. Francesco: 56 valves (BD 709), Carrabbati: 5 valves (BD 710), Castellace: 1 valve (BD 711), Gallina: 9 valves (BD 712), Le Castella: 150 valves (BD 713), Monasterace: 1 valve (BD 714), Musalà: 20 valves (BD 715), Pecoraro: 28 valves (BD 716), Petti di Carrubbare: 3 valves (BD 717), Pezzo: 134 valves (BD 718), S. Maria di Catanzaro: 11 valves (BD 719), Stalettì: 6 valves (BD 720), Terreti: 15 valves (BD 721), Torrente Boscaino: 2 valves (BD 722); Sicily: Capo Milazzo: 67 valves (BD 723), Grammichele: 1 valve (BD 724), Menfi: 3 valves (BD 725), Messina: 3 valves (BD 726), Salice: 7 valves (BD 727), Selinunte Casa Catarinicchia: 5 valves (BD 728), Selinunte Casa Parrino: 2 valves (BD 729). Greece: Kyllini: 20 valves (BD 730, DGUP). Maximum width of the valves: 6.7 / 8.3 / 6.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5D4EA60FADFAD56A9495E8.taxon	description	Description. Head valve semicircular, posterior margin markedly V-shaped, front slope straight. Intermediate valves broadly rectangular (W / L = 2.20 – 3.19), moderately elevated (H / W = 0.30 – 0.47), semicarinate in anterior profile, anterior margin evenly to weakly convex, side margins truncated, posterior margin straight but for the small, sharp-pointed apex, lateral areas raised, clearly separated from CA, sometimes by a clearly visible step. Tail valve semicircular (W / L = 1.70 – 2.13), anterior margin slightly convex, mucro subcentral, antemucronal slope straight to slightly convex, postmucronal slope straight to slightly concave. HV, LA and PMA smooth, concentric growth lines vaguely indicated. CA and AMA sculptured with up to 15 longitudinal folds at both sides of rather broadly triangular smooth jugum, innermost folds narrow, not reaching valve’s front margin. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses wide, rounded, connected across narrow, shallow sinus by a short, dentate jugal plate, slit formula 8 – 10 / 1 / 9 – 11, slits shallow, inequidistant, slit rays hardly or not indicated, teeth short, finely pectinate, eaves narrow, minutely porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5D4EA60FADFAD56A9495E8.taxon	discussion	Remarks. Rhyssoplax corallina (Risso, 1826) is a common extant species in the Mediterranean Sea whose fossil record encompasses the lower Miocene to the Pleistocene of the Mediterranean, northeastern Atlantic and Paratethys areas. Sacco (1897) did not report Chiton corallinus, and placed valves from Neogene deposits, with HV, LA and PMA smooth, as Chiton olivaceus var. plioparva.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5D4EA60FADFAD56A9495E8.taxon	description	Dell’Angelo et al. (2018 a) attributed tentatively to Rhyssoplax corallina two valves from the upper Oligocene (Chattian) of Saint-Paul-les-Dax (France, Aquitaine Basin). Given the relatively poor preservation state of the material and geological antiquity, we consider unlikley its co-specificity with the extant taxon (see under “ species of problematic assignment ”). The material examined displays a great variability in both the shape of the valves and the sculpture of the tegmentum, as reported by the material from the Pliocene of Liguria (Dell’Angelo et al. 2013) and the Miocene of N. Italy (Dell’Angelo et al. 2016). An example of the variability of the tail valves is shown in the three tail valves figured in lateral view (Figs 89 L, 89 N and 89 P), regarding the position of the mucro (subcentral in Figs 89 L and 89 P) and the antemucronal and postmucronal slopes. Furthermore, the sculpture of the central and antemucronal areas in Rhyssoplax corallina is not formed by true longitudinal ribs, but rather by longitudinal small folds (Figs 89 F, 89 I), as is also the case in R. olivacea. Comparisons. Rhyssoplax corallina is very similar to R. assurrecta Dell’Angelo et al., 2018 b (see above).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF5D4EA60FADFAD56A9495E8.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Aquitanian / Burdigalian): Aquitaine Basin, France: Carrière Vives, Coupe du fossé près de La Solitude, Lahitet, Maïnot (Dell’Angelo et al. 2018 a); Proto – Mediterranean Sea (Burdigalian): N. Italy: Valle Ceppi, Sciolze (Dell’Angelo et al. 2016). Middle Miocene: northeastern Atlantic (Serravallian): Aquitaine Basin, France: Orthez, Sallespisse (Dell’Angelo et al. 2018 a); Proto – Mediterranean Sea (Langhian): Po Basin, N. Italy: Albugnano (Dell’Angelo et al. 2016); Central Paratethys (Langhian – Serravallian): Slovakia: Devínska Nová Ves, Rohožník, Dubová (Ruman & Hudáčková 2015), Czech Republich: Knínice, Rudoltice, Zidlochovice (Procházka, 1895, 1900; Šulc 1934), Poland: Korytnica, Lychów, Nawodzice, Rybnica, Węglin, Weglinek (Bałuk 1971, 1984; Macioszczyk 1988; Studencka & Studencki 1988), Austria: Grund, Steinabrunn (Šulc 1934; Kroh 2003), Hungary: Bánd, Borsodbóta, Devecser, Letkés, Sámsonháza, Várpalota (Dulai 2005, 2025 a, 2025 b; Dulai & Katona 2024; Schwabe & Dulai 2024; this study), Romania: Bujtur, Costej, Lăpugiu de Sus (Rado 1969; Dell’Angelo et al. 2007 a: this study); Eastern Paratethys: Ukraine: Varovtsi, Zalistsi (Studencka & Dulai 2010; this study). Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place, Sceaux d’Anjou, La Presselière, Beugnon (Dell’Angelo et al. 2018 b); northeastern Atlantic (Messinian?), Ligerian Basin, France: Moulin Pochas (Dell’Angelo et al. 2018 a); Proto – Mediterranean Sea (Tortonian – Messinian): Po Basin, N. Italy: Villa Monti, Rio di Bocca d’Asino, Vargo, Vigoleno, Montegibbio, Borelli (Laghi 1977; Dell’Angelo et al. 1999, 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: many localities (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013). Pliocene: northeastern Atlantic, Mondego Basin, Portugal (MPMU 1); Vale de Freixo (Dell’Angelo & Silva 2003; Dell’Angelo et al. 2022); western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: many localities in Piedmont, Emilia-Romagna, Tuscany, Latium, Sicily (Laghi 1977; Dell’Angelo et al. 2001 a; Chirli 2004; this study). Upper Pliocene to upper Pleistocene: central Mediterranean, France: Bosq d’Aubigny (Dell’Angelo et al. 2018 b), Greece: Rhodes Island (Koskeridou et al. 2009). Pleistocene: North Atlantic: Netherlands (Strack 2010); central Mediterranean, Italy: many localities in Tuscany, Puglia, Calabria, Sicily (Sabelli & Taviani 1979; Dell’Angelo et al. 2001 a; this study); Greece: Kyllini (Garilli et al. 2005). Recent: Atlantic Ocean: Spain (Carmona Zalvide et al. 2000), Portugal (Consolado Macedo et al. 1999); Morocco (Pallary 1920); Ampère and Gettysburg Seamounts (Beck et al. 2006). Mediterranean Sea: Spain: Alboran isl. (Salas & Luque 1986); France (Marion 1883; Pruvot 1897); Italy: many localities (Monterosato 1879; Dell’Angelo & Smriglio 1999; Vazzana 2010; Trono & Macrì 2013); Croatia: (Leloup & Volz 1938; Dell’Angelo & Zavodnik 2004); Greece and Aegean Sea Islands (Strack 1988, 1990; Zenetos & Van Aartsen 1995; Koukouras & Karachle 2005); Turkey (Ozturk et al. 2014); Algeria (Mars 1957); Tunisia (Kaas 1989; Cecalupo et al. 2008); Israel (Barash & Danin 1977); Lebanon (Crocetta et al. 2014); Marmara Sea (Öztürk et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF594EA70FADFEB4681196BC.taxon	description	Fig. 90	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF594EA70FADFEB4681196BC.taxon	materials_examined	Type material. Holotype MZB 11623 (intermediate valve) (Dell’Angelo et al. 2001 a). Paratypes: MZB 12692 (head valve), MZB 12695 (tail valve), MCZR (3 valves), Civico Museo Archeologico e di Scienze Naturali di Alba, n ° reg. G- 1125 (4 valves), and private collections (12 valves): BD FX 18 A / C 12 (5 valves, Figs 90 D – F), MF (5 valves), LD (1 valve), VB F 1001 a (1 valve). Type locality. Riparbella (Tuscany, Italy). Type stage. Lower Pleistocene. Material examined. Pliocene Spain: El Papiol: 63 valves (BD 731). Italy: Piedmont: Vintebbio: 135 valves (BD 732); Tuscany: Cetona “ Palazzo Tosoni ”: 6 valves (BD 733); Colle Val d’Elsa: 42 valves (BD 734); Orciano Pisano: 33 valves (BD 735); Pietrafitta: 200 valves (BD 736, Figs 90 G – H); Pietrafitta Melograni: 89 valves (BD 737); Poggio alla Fame: 38 valves (BD 738, Figs 90 J – L). Pleistocene: Italy: Tuscany: Fauglia: 68 valves (BD 739), Riparbella: type material plus 336 valves (BD 740, Figs 90 A – C, 90 I, MF); Calabria: Le Castella: 11 valves (BD 741). Maximum width of the valves: 2.3 / 2.5 / 3.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF594EA70FADFEB4681196BC.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight. Intermediate valves broadly rectangular (W / L = 1.96 – 2.19), elevated (H / W = 0.50 – 0.65), rounded to semicarinate in anterior profile, anterior margin straight to slightly convex, side margins truncated, posterior margin straight but for the small, sharp-pointed apex, lateral areas raised, clearly separated from CA. Tail valve semicircular (W / L = 1.57 – 1.65), anterior margin slightly convex, mucro in slightly anterior position, antemucronal slope straight, postmucronal slope slightly concave. HV, LA and PMA smooth, concentric growth lines vaguely indicated. CA and AMA sculptured with 5 - 8 longitudinal folds at both sides of smooth jugum, all folds reach up to anterior margin, with exception of most central one (or last two) on each side. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses wide, subtrapezoidal, connected across narrow, shallow sinus by a short, dentate jugal plate, slit formula 8 – 10 / 1 / 11 – 14, slits inequidistant, slit rays hardly or not indicated, teeth short, finely pectinate, eaves narrow, minutely porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF594EA70FADFEB4681196BC.taxon	discussion	Remarks. Rhyssoplax etrusca (Dell’Angelo & Forli, 1995) is only known from Pliocene and Pleistocene of Tuscany, very frequent at Riparbella, where is the most abundant chiton. Comparisons. Rhyssoplax etrusca is very similar to R. phaseolina (Monterosato, 1879), from which it differs mainly by the more elevated intermediate valves, H / W = 0.50 – 0.65 vs. 0.40 – 0.50 in R. phaseolina, LA raised, clearly separated from CA, and almost all the longitudinal folds on CA reaching the anterior margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF594EA70FADFEB4681196BC.taxon	distribution	Distribution. Pliocene: western Mediterranean, Spain: El Papiol (this study); central Mediterranean, Italy: Cetona, Colle Val d’Elsa, Orciano Pisano, Pietrafitta, Pietrafitta Melograni, Poggio alla Fame, Vintebbio (Dell’Angelo et al. 2001 a; Chirli 2004; this study). Pleistocene: central Mediterranean, Italy: Fauglia, Le Castella, Riparbella (Dell’Angelo & Forli, 1995 a; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	description	Fig. 91	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	materials_examined	Type material. Holotype MSNG 62661, tail valve, width 4.4 mm (Figs 91 J – L). Paratype 1: MSNG 62662, head valve, width 2.8 mm (Figs 91 A – B). Paratype 2: MSNG 62663, intermediate valve, width 3.5 mm (Figs 91 E – G). Paratype 3: MNHN. F. A 98476, intermediate valve, width 5.2 mm (Figs 91 C – D). Paratype 4: MNHN. F. A 98477, tail valve, width 4.2 mm (Figs 91 H – I). Paratype 5: SMF 380821, intermediate valve, width 5.8 mm. Paratype 6: SMF 380822, tail valve, width 4.2 mm). Type locality. Altavilla Milicia (Sicily, Italy). Type stage. Pliocene-Pleistocene (upper Piacenzian to lower Gelasian) (Dominici et al. 2020).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	etymology	Etymology. The specific name honors Vittorio Garilli, for his contribution to the study of Recent and fossil mollusks of the Mediterranean basin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	materials_examined	Material examined. Upper Pliocene: Italy: Altavilla: type material plus 37 valves (AG, AR, BD 742). Maximum width of the valves: 7.3 – 6 – 6.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	diagnosis	Diagnosis. Head valve semi-oval, intermediate valves broadly rectangular, moderately elevated, carinate, apex small, sharp-pointed, tail valve more or less triangular, mucro in anterior position. Tegmentum smooth in HV, LA, PMA, sculptured with up to 12 longitudinal folds at both sides of smooth jugum, aesthetes very dense. Articulamentum with apophyses wide, rounded, connected by a short, dentate jugal plate, teeth pectinate, slit formula: 9 / 1 / 9 – 12.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	description	Description. Head valve semi-oval, front slope straight, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.76 – 2.94), moderately elevated (H / W = 0.35 – 0.47), carinate in anterior profile, anterior margin straight between apophyses, side margins truncated, posterior margin straight but for the small, sharp-pointed apex, lateral areas somewhat raised, clearly defined. Tail valve more or less triangular, W / L = 1.76 – 2.10, anterior margin straight between apophyses, mucro in anterior position, antemucronal slope straight to slightly convex, postmucronal slope slightly concave. HV, LA and PMA smooth, concentric growth lines well marked. CA and AMA sculptured with up to 12 longitudinal folds at both sides of smooth jugum, innermost folds narrow, not reaching valve’s front margin. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses wide, rounded, connected across narrow, shallow sinus by a short, dentate jugal plate, slit formula: 9 / 1 / 9 – 12, slits shallow, inequidistant, slit rays finely indicated, teeth short, pectinate, eaves narrow, porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	discussion	Remarks. The known fossil record of Rhyssoplax garillii sp. nov. is limited thus far to the Pliocene of Altavilla Milicia, in Sicily. The material examined belongs to the group of Rhyssoplax spp. characterized by the smoothness of HV, LA and PMA. These features are shared by Rhyssoplax corallina (Risso, 1826), one of the most common species of the Italian Pliocene, R. phaseolina (Monterosato, 1879), R. etrusca (Dell’Angelo & Forli, 1995), R. assurrecta Dell’Angelo et al., 2018 and R. capecchii (Chirli, 2004). However, the valves from Altavilla Milicia differ from R. corallina for a series of fairly constant characters (constants within the broad range of variability known for R. corallina), and such as to justify the establishment of a new species. Rhyssoplax garillii sp. nov. has already been reported by Seguenza (1876) as “ Chiton miocenicus Michelotti, 1847 Var. ” with a short diagnosis: “ Questa forma pliocenica differisce dal tipo della Superga per le linee d’accrescimento impresse e meglio distinte ”. Seguenza underlined the presence of marked growth lines as a distinctive character from C. miocenicus, and his short description was most probably based on material conspecific with the present one. Comparisons. Rhyssoplax garillii sp. nov. differs from R. corallina (Risso, 1826) by the more or less triangular shape of the tail valves (more semicircular in R. corallina), the mucro in anterior position (subcentral in R. corallina), the anterior profile of the intermediate valves decidedly carinate (semicarinate and tending to be more rounded in R. corallina), the presence of concentric growth lines well marked (vaguely indicated in R. corallina).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF584EA80FADFC406A919234.taxon	distribution	Distribution. Upper Pliocene: central Mediterranean, Italy: Altavilla Milicia (Dell’Angelo et al. 2012; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF574EAA0FADF8C96E2892CA.taxon	description	Fig. 92	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF574EAA0FADF8C96E2892CA.taxon	materials_examined	Type material. MGR (fide Sacco 1897: pl. 7, fig. 11 “ es. tip. fig. ”). Type locality. The Turin hills (Piedmont, Italy). Type stage. Lower Miocene. Material examined. Lower Miocene: Italy (Burdigalian): Sciolze: 2 valves (PG), Valle Ceppi: 23 valves (PG). Middle Miocene: Italy (Langhian): Albugnano: 43 valves (MZB 32079, PG). Upper Miocene: Italy (Tortonian): Borelli: 406 valves (BD 743, MGPT PU 135296 - 135298, Figs 92 A – B, 92 L), MZB 32081, PG), Moncucco Torinese: 1 valve (PG), Montegibbio: 78 valves (BD 744, MZB 32080, Fig. 92 E, LB), Rio di Bocca d’Asino: 149 valves (BD 745, PG), Sardigliano: 2 valves (BD 746), Vargo: 1 valve (PG), Villa Monti: 11 valves (BD 747, PG). Lower Pliocene: Italy: Liguria: Borzoli: 396 valves (BD 748), Bussana: 4 valves (BD 749), Garlenda: 5 valves (MP), Rio Sant’Antonino: 332 valves (BD 750, MP, PG, MZB 45743 - 45746, Figs 92 F – H), Rio Torsero: 1 valve (BD 751). Pliocene: Spain: Estepona: 630 valves (BD 752, Figs 92 I – K); Italy: Piedmont: San Damiano: 1 valve (BD 753); Tuscany: Guistrigona: 1 valve (BD 754), Montenero: 163 valves (BD 755, Figs 92 C – D), Villa Banfi: 4 valves (BD 756). Maximum width of the valves: 14.5 / 23 / 17.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF574EAA0FADF8C96E2892CA.taxon	description	Description. Head valve semicircular, front slope faintly convex, posterior margin widely V-shaped, with minute notch in the middle. Intermediate valves broadly rectangular (W / L = 2.33 – 3.33), moderately elevated (dorsal elevation 0.24 – 0.43), carinate in anterior profile, anterior margin straight to slightly convex, side margins truncated to weakly convex, posterior margin slightly concave at both sides of minute apex. Tail valve semicircular (W / L = 1.58 – 1.95), anterior margin convex, mucro subcentral, not prominent, antemucronal slope slightly convex, postmucronal slope almost straight. HV, LA and PMA sculptured with rather coarse, flattish radiating ribs (HV 30 – 45, LA 5 – 11), often bifurcating, more irregular, crossed by concentric growth lines. CA and AMA sculptured with 10 – 35 longitudinal ribs at both sides of the smooth, more or less triangular jugum, all reaching the anterior margin. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses wide, rounded, tending to trapezoidal in tail valve, jugal sinus narrow, provided with a concave, dentate plate, teeth short, minutely denticulate, slit formula 8 – 10 / 1 / 13 – 14, slits narrow, slit rays very finely indicated, eaves narrow, porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF574EAA0FADF8C96E2892CA.taxon	discussion	Remarks. Rhyssoplax miocenica (Michelotti, 1847) closely resembles R. olivacea (Spengler, 1797), and was considered in the past either a synonym of the latter or a distinct species. It was described from the Miocene of the Turin hills; one intermediate valve was well figured (Michelotti 1847: pl. 16, fig. 7). Many subsequent authors considered the species a synonym of R. olivacea (e. g., Laghi 1977; Laghi & Russo 1978). Others considered, on the contrary, R. miocenica a valid species (e. g., Monterosato 1879); Sacco 1897). Gymnoplax orbignyi de Rochebrune, 1882 was originally proposed as “ nomen substitutivum ” for Chiton subcajetanus d’Orbigny, 1852, based on chiton valves from the Turin hills. A syntype of Gymnoplax orbignyi is preserved at MNHN (Dell’Angelo et al. 2015 a: pl. 2, figs 14 – 17), and examination of this material made it possible to consider Gymnoplax orbignyi a synonym of Rhyssoplax miocenica (Dell’Angelo et al. 2015 a). It is difficult to evaluate statistically the morphological characteristics of the two species, because of the insufficient number of complete valves of R. miocenica to examine. An attempt has been made with intermediate valves of Rhyssoplax olivacea [56 valves, from Miocene (4), Pliocene (2), Pleistocene (24) and living (26)] and R. miocenica [22 valves, from Miocene (6) and Pliocene (16)]. Some parameters were measured on each valve (width, length and height), and the numbers of ribs (or folds) on central and lateral areas were counted. The relationship between the valve’s width and the width / length and height / width ratios are showed in Tables 14 / 1 and 14 / 2. This study allowed to confirm R. miocenica as a distinct species, different from R. olivacea, and to better characterize these two species. Comparisons. This species is closest to Rhyssoplax olivacea (Spengler, 1797), from which it differs mainly for the sculpture of CA and AMA formed by real longitudinal ribs (Fig. 92 D, not small folds as in R. olivacea), and for several other characters, e. g., the greater number of longitudinal ribs on CA (10 – 35 on each side, vs. 6 – 16 folds in R. olivacea), LA (5 – 10 vs. 2 – 6 in R. olivacea), and the different shape of tail valves, with the antemucronal slope slightly convex (more convex in R. olivacea) and the postmucronal slope almost straight (mainly concave close to the mucro in R. olivacea).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF574EAA0FADF8C96E2892CA.taxon	distribution	Distribution. Lower Miocene: Proto-Mediterranean Sea (Burdigalian): N. Italy: Sciolze, Valle Ceppi (Michelotti 1847; Dell’Angelo et al. 2016). Middle Miocene: Proto-Mediterranean Sea (Langhian): Po Basin: Albugnano (Dell’Angelo et al. 2016). Upper Miocene: Proto-Mediterranean Sea (Tortonian-Messinian): Po Basin, N Italy: Borelli, Moncucco Torinese, Montegibbio, Rio di Bocca d’Asino, Sardigliano, Vargo, Villa Monti (Laghi 1977; Dell’Angelo et al. 1999, 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Garlenda, Rio S. Antonino, Rio Torsero (Dell’Angelo et al. 2001 a, 2013; Sosso & Dell’Angelo 2010). Pliocene: western Mediterranean, Estepona Basin, Spain (Dell’Angelo et al. 2004); central Mediterranean, Italy: Piedmont: San Damiano; Tuscany: Guistrigona, Montenero, Villa Banfi (Chirli 2004; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF544EAF0FADF97C6B6B91B1.taxon	description	Fig. 93	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF544EAF0FADF97C6B6B91B1.taxon	materials_examined	Type material. Presumably lost, the type series of Chiton olivaceus is no longer present in the collection of the Zoological Museum of the University of Copenhagen (fide Kaas & Knudsen 1992: 60). Type locality. Off the coast of North Africa. Material examined. Middle Miocene: Italy (Langhian): Albugnano: 2 valves (PG); Central Paratethys (Langhian-Serravallian): Austria: Gainfarn: 2 valves (BD 757), Niederkreuzstetten: 1 valve (NHMW 1859 / 0045 / 0077), Pötzleinsdorf: 40 valves (NHMW 1859 / 0027 / 0063, 1861 / 0028 / 0078, 2010 / 0256 / 0006); Romania: Bujtur: 3 valves (NHMW 1863 / 0015 / 0116), Kostej: 12 valves (BD 758, NHMW 1869 / 0001 / 0796, 1870 / 0049 / 0126, 2010 / 0256 / 0011), Lăpugiu de Sus: 55 valves (BD 759, NHMW 1854 / 0035 / 0407, 1865 / 0001 / 0273, 1868 / 0001 / 0633, 1876 A / 0011 / 0175); Czech Republic: Porzteich: 23 valves (NHMW 1871 / 0010 / 0210), Rousinov: 1 valve (NHMW 1861 / 0011 / 0409); Hungary: Bánd: 54 valves (BD 760, Fig. 93 E – G), Letkés: 17 valves (BD 761), Màrkhàza: 1 valve (BD 762); Eastern Paratethys: Ukraine: Horodok: 17 valves (BD 763), Varovtsi: 65 valves (BD 764, Fig. 93 B – D), Velyka Levada: 1 valve (BD 765). Upper Miocene: Italy: Borelli: 4 valves (MGPT PU 135049, PG); Montegibbio: 27 valves (BD 766, MZB 32070); Rio di Bocca d’Asino: 196 valves (BD 767, Fig. 93 M – O, PG, MZB 32065 - 32069, Figs 93 A, 93 H, 93 P); S. Agata Fossili: 3 valves (BD 768); Vigoleno: 7 valves (BD 769); Villa Monti: 16 valves (BD 770). Lower Pliocene: Italy: Liguria: Borzoli: 10 valves (BD 771); Bussana: 8 valves (BD 772, MZB 45728 - 45729, Fig. 93 I – J); Caranchi: 3 valves (MP); Genova Sestri: 3 valves (BD 773); Rio Torsero: 2 valves (BD 774). Pliocene: Italy: Piedmont: Baldichieri: 1 valve (BD 775), San Damiano: 12 valves (BD 776), Villalvernia: 4 valves (BD 777), Vintebbio: 3 valves (BD 778); Tuscany: Castiglioncello del Trinoro: 3 valves (BD 779), Grotte di Pagnana: 4 valves (BD 780); Emilia-Romagna: Gagliardella “ Tagliata ”: 1 valve (BD 781), Marano sul Panaro: 1 valve (BD 782), Torrente Stirone: 2 valves (BD 783); Latium: Montelibretti: 1 valve (BD 784): Sicily: Trappeto: 4 valves (BD 785). Pleistocene: Greece: Kyllini: 48 valves (BD 786, DGUP, Fig. 93 K – L), Loutraki: 33 valves (BD 787); France: Corsica, Patrimonio: 1 valve (BD 788); Italy: Emilia-Romagna: Torrente Guerro: 1 valve (BD 789), Torrente Stirone: 20 valves (BD 790); Tuscany: Cisternino: 13 valves (BD 791), Fauglia: 1 valve (BD 792), Riparbella: 12 valves (BD 793); Campania: Marina di Camerota: 5 valves (BD 794); Puglia: Gallipoli: 3 valves (BD 795), Mar Piccolo: 13 valves (BD 796), Punta Penne: 7 valves (BD 797); Calabria: Archi S. Francesco: 78 valves (BD 798), Bovetto: 1 valve (BD 799), Carrabbati: 3 valves (BD 800), Gallina: 2 valves (BD 801), Le Castella: 432 valves (BD 802), Musalà: 11 valves (BD 803), Pecoraro: 5 valves (BD 804), Petti di Carrubbare: 3 valves (BD 805), Pezzo: 58 valves (BD 806), S. Maria di Catanzaro: 4 valves (BD 807), San Procopio: 5 valves (BD 808), Saracinello: 1 valve (BD 809), Stalettì: 15 valves (BD 810), Torrente Boscaino: 1 valve (BD 811), Vallone Catrica: 1 valve (BD 812), Vito Superiore: 2 valves (BD 813); Sicily: Capo Milazzo: 1 valve (BD 814), Ficarazzi: 1 valve (BD 815), Grammichele: 8 valves (BD 816). Maximum width of the valves: 5 / 12 / 10.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF544EAF0FADF97C6B6B91B1.taxon	description	Description. Head valve semicircular, front slope faintly convex, posterior margin widely V-shaped, with minute notch in the middle. Intermediate valves broadly rectangular (W / L = 2.09 – 3.19), rather elevated (dorsal elevation 0.33 – 0.55), carinated in anterior profile, anterior margin slightly forwardly produced in small central part, side parts somewhat wavy, side margins truncated to weakly convex, posterior margin slightly concave at both sides of minute apex. Tail valve semicircular (W / L = 1.70 – 2.26), anterior margin convex, mucro subcentral, prominent, antemucronal slope slightly convex, postmucronal slope almost straight to slightly concave. HV, LA and PMA sculptured with rather coarse, flattish radiating ribs (HV 30 – 40, LA 2 – 6, PMA 25 – 35), often bifurcating, interstices narrower, crossed by several concentric growth lines. CA and AMA sculptured with 14 – 32 longitudinal folds, jugal area smooth, somewhat with traces of longitudinal folds near apex, innermost folds narrow, close set, not reaching front margin, others gradually becoming broader, more widely spaced and covering entire length of the valve. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses wide, rounded, tending to trapezoidal in tail valve, jugal sinus narrow, about half of the valve’s width, deep, provided with a concave, dentate plate, teeth short, minutely denticulate, slit formula 8 – 10 / 1 / 9 – 14, slits narrow, slit rays very finely indicated, eaves narrow, porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF544EAF0FADF97C6B6B91B1.taxon	discussion	Remarks. Rhyssoplax olivacea (Spengler, 1797) is the most common and best-known Mediterranean species; its remarkable variability led to the description of several species, currently considered synonyms of R. olivacea (Dell’Angelo & Smriglio 1999; Kaas et al. 2006). Regarding fossil valves, their morphological variability is even stronger, resulting in the description of several taxa, e. g., Chiton zibinicus Doderlein, 1864 from the Tortonian of Montegibbio, and two species from the Paratethys, Gymnoplax bohemicus de Rochebrune, 1882 and Chiton jüttneri Šulc, 1934. Chiton zibinicus Doderlein, 1864 was described with the diagnosis: “ Ch. miocenici Micht. prox. sed distinctus ”, without illustrations. Sacco (1897) considered this species as a variety of C. olivaceus, and figured a tail valve (pl. 7, figs 6 – 7). We follow Sacco (1897), Laghi (1977) and Dell’Angelo et al. (2016) in considering Chiton zibinicus as a synonym of Rhyssoplax olivacea.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF544EAF0FADF97C6B6B91B1.taxon	materials_examined	The material examined show great variability in the sculpture of the tegmentum, the longitudinal folds on PA may be parallel to each other (Figs 93 B, 93 I, 93 K) or obliquely arranged towards the jugal region (Fig. 93 H), which can be smooth for a large part (Figs 93 E, 93 K) or almost completely covered by folds (Fig. 93 M). The longitudinal folds on PA normally all reaching the anterior margin, but in some cases the folds near the jugum stop just short of it (Figs 93 H – I). Furthermore, the number of longitudinal folds and radial ribs is variable; we have recorded 14 – 32 longitudinal folds and 2 – 6 radial ribs in the intermediate valves examined. It should be noted that the sculpture of CA and AMA is not formed by real longitudinal ribs, as in many Rhyssoplax spp. (e. g., R. miocenica), but rather by the spaces between small folds on the tegmentum, as already reported by Laghi (1977: p. 110 “ aree pleurali incise da solchi longitudinali che in realtà sono gli spazi tra piccole pieghe del tegmentum rovesciate verso il perinoto ”), and as can be seen in the figured valve (Figs 93 J, 93 L). Some valves show a well-defined dental plate in the jugal sinus, as the figured valve (Figs 93 E – G) from the Paratethys. The sculpture of the dorsal surface has been well highlighted by Fischer & Renner (1979). Comparisons. The more similar species is Rhyssoplax miocenica (Michelotti, 1847), and the differences with this species are discussed above.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF544EAF0FADF97C6B6B91B1.taxon	distribution	Distribution. Middle Miocene: Mediterranean Sea (Langhian): France: La Chausserie (Dollfus 1920); Italy: Po Basin: Albugnano (Dell’Angelo et al. 2016); Central Paratethys (Langhian-Serravallian): Austria: Gainfarn, Niederkreuzstetten, Pötzleinsdorf (Šulc, 1934); Czech Republic: Borač, Knínice, Porzteich, Rousinov, Rudoltice (Šulc 1934); Slovakia: Rohožník (Ruman & Hudácková 2015); Poland: Lychów, Niskowa, Węglin, Weglinek (Skoczylasówna 1930; Bałuk, 1965; Macioszczyk, 1988); Hungary: Bánd, Devecser, Letkés, Màrkhàza (Dulai 2005, 2025 a, 2025 b; this study), Romania: Băseşti, Bujtur, Coştei, Lăpugiu de Sus (Šulc 1934; Zilch 1934; Marinescu, 1964; Dell’Angelo et al. 2007 a); Eastern Paratethys: Ukraine: Horodok, Varovtsi, Velyka Levada (Studencka & Dulai, 2010; this study). Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Borelli, Montegibbio, Rio di Bocca d’Asino, S. Agata Fossili, Vigoleno, Villa Monti (Laghi 1977; Dell’Angelo et al. 1999, 2016). Lower Pliocene: northeastern Atlantic: U. K. (Reid 1890, 1892); central Mediterranean, Italy: Liguria: many localities (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013). Pliocene: central Mediterranean, Italy: many localities in Piedmont, Emilia-Romagna, Tuscany, Latium, Sicily (Dell’Angelo et al. 2001 a, 2013; Sosso & Dell’Angelo 2010; this study); France: Biot (B. D. D. 1882). Upper Pliocene to upper Pleistocene: Greece: Rhodes Island (Koskeridou et al. 2009). Lower Pleistocene: central Mediterranean, Italy: Torrente Stirone (Sabelli & Taviani 1979). Pleistocene: central Mediterranean: Greece and Cyprus (Garilli et al. 2005); France: Corsica, Patrimonio (this paper); Italy: many localities in Emilia-Romagna, Tuscany, Campania, Puglia, Calabria, Sicily (Dell’Angelo et al. 2001 a; this paper), Tunisia: Monastir (Castany et al. 1956), Algeria: Arzeu (De Lamothe 1911). Recent: Atlantic Ocean, Spain (Carmona Zalvide et al. 2000), Portugal (Consolado Macedo et al. 1999), Morocco (Pallary 1920) and Berlengas Arch. (Pisani Burnay 1986); Mediterranean Sea. Spain (Salas & Luque 1986; Giribet & Penas 1997), France (B. D. D 1982), Italy: many localities (Monterosato 1879; Dell’Angelo & Smriglio 1999; Kaas et al. 2006; Trono 2006); Croatia: Dell’Angelo & Zavodnik 2004; Greece and Aegean Sea Islands (Strack 1988, 1990; Koukouras & Karachle 2005); Malta (Mifsud et al. 1990); Turkey (Öztürk et al. 2014); Tunisia (Kaas 1989; Cecalupo et al. 2008); Israel (Barash & Danin 1977); Lebanon (Crocetta et al. 2014); Marmara Sea (Öztürk et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF504ED00FADFB4F681E9376.taxon	description	Fig. 94	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF504ED00FADFB4F681E9376.taxon	materials_examined	Type material. MCZR – M – 12674: 3 lots (1 spm from “ Palermo Acqua Santa ”; 1 spm from “ Golfo di Napoli ”; 2 spm and 5 valves from “ Palermo Acqua Santa ” (Appolloni et al. 2018). Type locality. Described from Palermo (Sicily, Italy) and Gulf of Napoli (Campania, Italy). Material examined. Pleistocene: Italy: Calabria: Gallina: 4 valves (AV, Figs 94 E – F, BD 817); Sicily: Ustica: 1 valve (MGUP 538, Figs 94 C – D). Recent: Mediterranean Sea: Italy: Ognina (BD 818), valves (Figs 94 A – B, 94 G – H). Maximum width of the valves: -- / 2.9 / 2.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF504ED00FADFB4F681E9376.taxon	description	Description. Head valve somewhat more than semicircular, posterior margin widely V-shaped, front slope weakly convex. Intermediate valves broadly rectangular (W / L = 2.43 – 2.66), moderately elevated (H / W = 0.40 – 0.50), rounded to semicarinate in anterior profile, anterior margin about straight in a wide central part, roundly sloping at the sides, posterior margin straight to slightly concave at both sides of prominent, sharply pointed apex, lateral areas little raised, not separated from CA. Tail valve semicircular (W / L = 1.62), anterior margin weakly angularly convex, mucro at anterior third, rather prominent, antemucronal slope almost straight, postmucronal slope straight to slightly concave. HV, LA and PMA smooth, concentric growth lines vaguely detectable. CA and AMA sculptured with 5 – 8 longitudinal folds on each side, of which only first two or three reach up to anterior margin, others cover a half or a third of length. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses wide, triangular with rounded top, connected across shallow sinus by short, finely denticulate jugal plate, insertion plates short, slit formula 8 – 14 / 1 / 8 – 9, slit rays hardly discernible, teeth rather blunt, finely grooved on the outside, eaves very narrow, finely porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF504ED00FADFB4F681E9376.taxon	discussion	Remarks. The fossil record of Rhyssoplax phaseolina (Monterosato, 1879) known to date is limited to the Pleistocene of southern Italy; the valves here figured, from Ustica island (Sicily) and Gallina (Apulia) represent the first record of this taxonl. The species was formally described by Monterosato in 1879, although the taxon got mentioned in previous works: in 1872, as a variety of Chiton rubicundus O. G. Costa (“ Var. = C. phaseolinus, mihi ms. ”); in 1875, still as a variety of C. rubicundus; and in 1878 a. Comparisons. Rhyssoplax phaseolina could be mistaken for R. corallina (Risso, 1826), from which it differs mainly by its smaller size, semicarinate profile of the intermediate valves, lateral areas little raised, not separated from the central area, and the mucro located more anteriorly. For the differences with Rhyssoplax etrusca see above.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF504ED00FADFB4F681E9376.taxon	distribution	Distribution. Pleistocene: central Mediterranean, S. Italy: Gallina, Ustica (this study). Recent: Atlantic Ocean: southwestern Spain: Cádiz (Carmona Zalvide et al. 2000). Mediterranean Sea: Spain (Capo de Gata, Algeciras), S. Italy (Marina di Camerota, Sicily, Lampedusa, Pantelleria), Malta, Cyprus (Dell’Angelo & Smriglio 1999; Kaas et al. 2006), Lebanon (Crocetta et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2F4ED20FADF90E6E3D95C4.taxon	description	Fig. 95	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2F4ED20FADF90E6E3D95C4.taxon	materials_examined	Type material. Holotype MPM, intermediate valve figured by Laghi (1984: pl. 1, fig. 6 a – b). Type locality. Serre di Rapolano (“ Il Campino ”), Siena (Tuscany, Italy). Type stage. Pliocene, “ sabbie gialle senesi ”. Material examined. Pliocene: Italy: Tuscany: Cetona: 1 valve (BD 819), Cisternino: 2 valves (BD 820), Linari: 1 valve (BD 821), Montaione: 1 valve (BD 822), Orciano Pisano: 2 valves (BD 823), Pietrafitta: 54 valves (BD 824), Pietrafitta Melograni: 48 valves (BD 825); Serre di Rapolano: 1500 + valves (BD 826, Figs 95 A – L), Villa Banfi: 2 valves (BD 827); Umbria: Castel Viscardo: 1 valve (BD 828), Corbara: 1 valve (BD 829), Fabro Scalo: 2 valves (BD 830). Maximum width of the valves: 15 / 20 / 14 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2F4ED20FADF90E6E3D95C4.taxon	description	Description. Head valve semicircular, front slope almost straight to faintly convex, posterior margin widely V-shaped. Intermediate valves broadly rectangular, elongated (W / L = 2.20 – 3.44), moderately elevated (H / W = 0.27 – 0.39), carinated in anterior profile, anterior margin straight to slightly convex, side margins slightly rounded, posterior margin almost straight, minute apex. Tail valve semicircular (W / L = 1.81 – 2.02), anterior margin almost straight, mucro subcentral, antemucronal slope straight, postmucronal slope almost straight to weakly concave. Tegmentum smooth, crossed by several well marked growth lines, sometimes very light flat ribs are present, radial on HV, LA and PMA, longitudinal on CA and AMA. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses rounded, tending to trapezoidal in tail valve, jugal sinus narrow, provided with a concave, dentate plate, teeth minutely denticulate, slit formula 11 / 1 / 12 (a second slit ray lies close to posterior margin on intermediate valves), slits narrow, slit rays weekly marked.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2F4ED20FADF90E6E3D95C4.taxon	discussion	Remarks. The fossil record of Rhyssoplax saeniensis (Laghi, 1984) is limited to the Pliocene of Italy; the species is very abundant at the site of its first discovery (Serre di Rapolano), much less frequent (a few valves) in the other Pliocene localities from Tuscany and Umbria. Despite the large size, it is very difficult to find complete valves, they are almost always incomplete or abraded (Fig. 95 I), with the tegmentum often densely bored by endobionts (Fig. 95 D). Comparisons. Laghi (1984) compared Rhyssoplax saeniensis with R. olivacea (Spengler, 1797), illustrating the main morphological differences between the two taxa (Laghi 1984: fig. 1); this Author also highlighted the different sculpture of R. olivacea (not formed by real longitudinal grooves / ribs more or less evident as in R. saeniensis, but rather by the spaces between small folds on the tegmentum).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2F4ED20FADF90E6E3D95C4.taxon	distribution	Distribution. Pliocene: central Mediterranean, Italy: Tuscany: Cetona, Cisternino, Montaione, Orciano Pisano, Pietrafitta, Pietrafitta Melograni, Serre di Rapolano, Linari, Villa Banfi (Laghi 1984; Dell’Angelo et al. 2001 a; Chirli 2004; this study); Umbria: Castel Viscardo, Corbara, Fabro Scalo (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2D4ED30FADFED8681494EC.taxon	description	Fig. 96	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2D4ED30FADFED8681494EC.taxon	materials_examined	Type material. Holotype: MZB 32084 (tail valve, width 2.5 mm, Figs 96 F – H). Paratypes: MZB 32082 (intermediate valve from Albugnano, Fig. 96 C); MZB 32083 (tail valve from Albugnano); MZB 32085 (head valve from Montegibbio, Figs 96 A – B). Type locality. Rio di Bocca d’Asino (Piedmont, Italy). Type stage. Miocene (Tortonian). Material examined. Middle Miocene: Italy: Albugnano: type material plus 1 valve (PG). Upper Miocene: Italy: Rio di Bocca d’Asino: type material plus 2 valves (BD 831); Montegibbio: 1 valve (BD 832). Lower Pliocene: Italy: Liguria: Borzoli: 4 valves (BD 833, MZB 45748, Fig. 96 E), Bussana: 2 valves (BD 834), Genova Sestri: 2 valves (BD 835, MZB 45749, Fig. 96 D). Maximum width of the valves: 4.6 / 3.6 / 3.9 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2D4ED30FADFED8681494EC.taxon	description	Description. Head valve semicircular. Intermediate valve broadly rectangular, anterior margin straight between apophyses, posterior margin little concave at both sides of well pronounced apex, lateral areas raised. Tail valve semicircular (W / L = 1.84 – 1.88), little elevated, small anterior mucro, antemucronal slope straight, postmucronal slope concave. HV, LA and PMA smooth, except some light radial ribs restricted to area near margins. CA and AMA smooth, except for 3 – 4 small ribs each side obliquely directed, near margins. Aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum scarcely visible, slit formula: 8 – 9 / 1 / 10.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2D4ED30FADFED8681494EC.taxon	discussion	Remarks. The fossil record of Rhyssoplax sulcomarginata (Dell’Angelo, Giuntelli, Sosso & Zunino, 2016) is limited to the Miocene and Pliocene of Italy; the material examined is represented by small and badly preserved valves. The description is therefore less complete than that of the other Rhyssoplax spp. Comparisons. The main differential character is the presence of 3 – 4 small ribs obliquely directed, in PA and AMA areas near the side margin, and of some light ribs only in a small part near the margins of HV, LA and PMA.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2D4ED30FADFED8681494EC.taxon	distribution	Distribution. Middle Miocene: Proto-Mediterranean Sea (Langhian): Po Basin: Albugnano (Dell’Angelo et al. 2016). Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Rio di Bocca d’Asino, Montegibbio (Dell’Angelo et al. 2016). Lower Pliocene: central Mediterranean, N Italy, Liguria: Borzoli, Bussana, Genova Sestri (Dell’Angelo et al. 2013).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2C4ED30FADFDB06BFB91DF.taxon	type_taxon	Type species. Chiton confossus Gould, 1846 (= Chiton lamellosus Quoy & Gaimard, 1835), by subsequent designation (Pilsbry 1893).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2C4ED30FADFDB06BFB91DF.taxon	distribution	Distribution. Lucilina is known from the Oligocene to the Recent, with a modern distribution in the Indo-Pacific Ocean. The fossil record is quite scant, and includes the Oligocene of France (Dell’Angelo et al. 2018 a), the Miocene and Pliocene of the W. Pacific islands (Ladd 1966; Schwabe et al. 2008), and the Pleistocene of the Red Sea (Dell’Angelo et al. 2020 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2C4ED30FADFDB06BFB91DF.taxon	discussion	Remarks. Following the suggestions of some authors (e. g., Schwabe et al. 2008; Schwabe & Pittman 2014), and according to WoRMS (2024), the genus Lucilina is here considered as a separate genus, not a subgenus of Tonicia Gray, 1847. It differs from the latter mainly by the position of the mucro in tail valve, subterminal in Lucilina and subcentral in Tonicia (Kaas et al. 2006). Moreover, the species of Tonicia are presently restricted to the southeastern Pacific region off south America, with one single species reported for the Caribbean; on the contrary, members of Lucilina are widespread throughout the entire tropical Indo-Pacific. The main characters of the species considered here are given in Tab. 16. personal observations).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2B4ED50FADFD306B5F95E8.taxon	description	Fig. 97	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2B4ED50FADFD306B5F95E8.taxon	materials_examined	Type material. Holotype: MHNBx 2017.10.1, tail valve, width 3.6 mm (Figs 97 A – D). Paratype: MZB 32136, tail valve. Type locality. Gaas, Espibos (France). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene: France: Gaas: type material plus 3 tail valves, maximum width 5 mm (AC, DA, PR).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2B4ED50FADFD306B5F95E8.taxon	description	Description. Head and intermediate valves unknown. Tail valve almost semicircular (W / L = 2.07 for the paratype), anterior margin about straight, mucro small, raised, in posterior position, antemucronal slope concave, postmucronal slope steep, straight to weakly convex. AMA sculptured with oval, elevated granules, jugal area eroded. Articulamentum with apophyses well developed, broad, jugal sinus narrow, ca. 8 – 9 teeth relatively long, solid, forwardly directed.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2B4ED50FADFD306B5F95E8.taxon	discussion	Remarks. This species is known only for five tail valves from the Oligocene (Rupelian) of Gaas (France); this record extended the distribution of the genus Lucilina to the lower Oligocene. This is the only species of Lucilina known for the European Cenozoic. Some species from the Eocene of France were attributed in the past to the genus Tonicia, which could be, in fact, possibly belonging to Lucilina. The only species from the European Cenozoic attributed with certainty to Tonicia is T. pannonica Szőts, 1953 from the middle Eocene of Gant, Hungary, characterized by a smooth tegmentum and the presence of regular rows of subcircular ocelli (Dell’Angelo et al. 2015: fig. 3).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2B4ED50FADFD306B5F95E8.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2018 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED50FADFEB468409794.taxon	type_taxon	Type species. Chaetopleura gaasi Cherns & Schwabe, 2019.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED50FADFEB468409794.taxon	distribution	Distribution. Spinochiton is known only from the Oligocene (Rupelian) of Gaas (France).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED50FADFEB468409794.taxon	discussion	Remarks. The name Spinochiton (and Spinochitonidae) originally introduced by Dell’Angelo et al. (2018 a) were not validly described in that publication (Philippe Bouchet, in litteris) because the designated type species Chaetopleura gaasi Cherns & Schwabe (2019), was not yet available at the time of publication of Spinochiton. Both taxa were subsequently validly described by Dell’Angelo et al. (2019 c). The main characters of the species considered here are given in Tab. 16.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED70FADFD6868B195E8.taxon	description	Fig. 98	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED70FADFD6868B195E8.taxon	materials_examined	Type material. Holotype ZSM Mol 20060779 A, tail valve. Paratype ZSM Mol 20060779 D, head valve. Type locality. Gaas (France). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene: France: Gaas: 60 valves (AC, BD 836, Figs 98 K – M, DA, JVC, MHNBx 2017.7.3, Figs 98 A – E, MHNBx 2017.10.2, Figs 98 F – J, MHNBx 2017.11.1, Figs 98 N – P). Maximum width of the valves: 6.2 / 8.6 / 5.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED70FADFD6868B195E8.taxon	description	Description. Head valve semicircular, elevated, apex slightly recurved, front slope concave, posterior margin V-shaped, notched in middle. Intermediate valve broadly rectangular (W / L = 2.41 – 2.53), carinate in anterior profile, elevated (H / W = 0.45 – 0.54), anterior margin convex with jugal part forwardly produced, side margins oblique, posterior margin straight, apex inconspicuous, lateral areas sligthly raised. Tail valve almost semicircular (W / L = 1.53 – 1.70) anterior margin about straight with jugal part forwardly produced, mucro not prominent, in central position, antemucronal slope slightly convex, postmucronal slope concave just underneath mucro. HV sculptured with 13 – 14 radial rows of conical spinous processes rounded at top (without any type of sculpture), rows starting from a small smooth area around semicircular notch with spinous processes initially small and growing and getting more elevate towards anterior margin, 6 – 7 spinous processes well developed for each row, and 2 – 3 obsolete ones near notch. Two rows of spinous processes are on two sides of posterior margin, surface between striae rough, with evidence of growth lines but without a well-defined sculpture, everywhere perforated by micraesthetes, but not on spinous processes. LA sculptured with 2 radial rows of conical spinous processes rounded at top, like in HV, obsolete near apex and growing towards side margins, one of them on posterior margin. PMA sculptured with conical spinous processes that seem irregularly arranged, not in radial rows as in HV. CA and AMA sculptured with longitudinal coarse ribs with from 2 to 6 very irregular oval pustules, getting larger and fused together towards JA, latter formed by a single longitudinal rib with larger pustules fused together to reach two adjacent ribs, interspaces with growth lines vaguely indicated, intersected by very small and fine striae of longitudinal granules, giving a reticulate appearance. Articulamentum strongly developed with apophyses expanded, rounded, connected by a jugal lamina with external slits in intermediate and tail valves; head valve with a large, elevated apical area, extending obliquely to articulamentum, traversed by a series of more or less parallel and columnar thickenings branching towards apex; intermediate valve with a smaller but similar apical area; teeth solid, finely striated on upper side, in head valve corresponding to radial rows of spines, except for two near posterior margin for each side, slit rays vaguely evidenced, eaves solid, slit formula 9 – 13 / 1 / 11.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED70FADFD6868B195E8.taxon	discussion	Remarks. Cherns & Schwabe published, firstly online (1 Nov 2017) the description of the new species Chaetopleura gaasi, based on 4 valves (1 head and 3 tail) from Gaas. The printed version appeared later (Cherns & Schwabe 2019). The same species was studied simultaneously by Dell’Angelo et al. (2018 a), on a greater number of valves (60), also including intermediate valves (not present in the Cherns & Schwabe material). Their description strongly completemented what reported by Cherns & Schwabe (2019), highlighting some important morphologically characters that induced Dell’Angelo et al. (2018 a) to define the new genus Spinochiton. The large apical area extending obliquely to the articulamentum (Figs 98 E, 98 I, 98 L – M) is a weird character, not present in chitons from the Cenozoic, where the apical area is normally little evident in head valve, and more pronounced but always narrow in intermediate ones. Comparisons To our knowledge, no living or fossil chitons share, the same characters of Spinochiton gaasi. A few species show the head valve character of the apex slightly recurved and front slope strongly concave, e. g., living species of the genus Nuttallochiton Plate, 1899 (Kaas & Van Belle 1987), and the fossil Oochiton halli Ashby, 1929 from the Miocene of Australia (Ashby 1929; Smith 1960).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF2A4ED70FADFD6868B195E8.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Cherns & Schwabe 2019; Dell’Angelo et al. 2018 a, 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED70FADFEFC6ED197CC.taxon	type_taxon	Type species. Chiton incisus Sowerby, 1841, by monotypy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED70FADFEFC6ED197CC.taxon	distribution	Distribution. Schizochiton is known from the Paleocene to Miocene and the Recent. Only two living species are known, Schizochiton incisus (Sowerby, 1841) from the Red Sea up to Philippines and Indonesia, and S. jousseaumei Dupuis, 1917 from the Gulf of Aden. The fossil record includes the Paleocene of Belgium and Ukraine (Makarenko 1976), the Eocene of Ukraine (Bielokrys 1999), the Oligocene (Rupelian) of France (Dell’Angelo et al. 2018 a), and the Miocene of W. Pacific Islands (Ladd 1966).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED70FADFEFC6ED197CC.taxon	discussion	Remarks. The main characters of the species here discussed are summarized in Tab. 16.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED80FADFCD06B5F91A3.taxon	description	Fig. 99	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED80FADFCD06B5F91A3.taxon	materials_examined	Type material. Holotype: MHNBx 2017.7.4, head valve, width 2.3 mm (Figs 99 A – D). Paratype: MHNBx 2017.7.5, intermediate valve (Figs 99 E – H). Type locality. Gaas, Lagouarde (France). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene: France: Gaas, Lagouarde: type material plus 2 head valves (AC). Maximum width of the valves: 5.2 / 4.4 / -- mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED80FADFCD06B5F91A3.taxon	description	Description. Tail valve unknown. Head valve semioval, front slope slightly convex, apex small and distinct. Intermediate valves polygonal, as long as wide (WT / LT = 1.05), semicarinate in anterior profile, elevated (H / W = 0.5), anterior margin straight in central part, almost straight and strongly slanting at pleurae, side margins short, posterior margin straight at both sides of moderately produced, bluntly pointed apex, lateral areas small, hardly or not raised. HV sculptured with six raised rows of large pustules growing outward toward anterior margin, with deep holes on top of each pustule, interspaces between pustule rows with many large and well separated oval granules, smaller near apex and growing outward anterior margin. LA sculptured with an elevated diagonal row of large pustules with deep holes on top, and a more obsolete second radial row of pustules at posterior margin. PA sculptured with large oval granules, well separated near diagonal rows of pustules between PA and LA, becoming united and forming short segments, acquiring vaguely zig-zag pattern near JA, triangular, longitudinally finely striated, without any other sculpture. Articulamentum with subrectangular apophyses, insertion plates long, slit formula 6 / 1 / --, slits shallow, corresponding to tegmental rows of pustules, slit rays indicated, teeth sharp, finely striated on the upper side.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED80FADFCD06B5F91A3.taxon	discussion	Remarks. The deep V-shaped caudal sinus in the tail valve, one of the main characters of the genus Schizochiton Gray, 1847 (Kaas et al. 2006: p. 49), is unfortunately not comparable in the studied specimens, lacking tail valves. However, the other characters are sufficient to attribute the valves examined to the genus Schizochiton. The valves are in some cases eroded, and, therefore, not characteristics are fully detectable in detail. The holes on the top of large pustules could be likely interpreted as eyepiece housing for the ocelli; this aspect is reported from living representatives of Schizochiton but are not, to the best of our knowledge, normally preserved in fossils species, as is the case of the specimens here figured. The holes seem deep, penetrating inside the valves. Comparisons. Bielokrys (1999) described three species of Schizochiton from the Eocene of Ukraine: S. parcus, S. carinatus and S. hirtus, different from S. tasteti (see Dell’Angelo et al. 2018 a for comparisons).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF284ED80FADFCD06B5F91A3.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2018 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF274EDB0FADFA126E7C94B4.taxon	type_taxon	Type species. Chiton marginatus Pennant, 1777 (= Chiton cinereus Linnaeus, 1767), by monotypy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF274EDB0FADFA126E7C94B4.taxon	distribution	Distribution. Lepidochitona is known from the Eocene to the Recent, with a present distribution in five areas: the Black and Mediterranean seas and the north-east Atlantic Ocean from Morocco to north Norway, the Red Sea and north Arabian Sea, off South Africa, the Caribbean Sea and adjacent waters of the Atlantic Ocean, eastern part of the Pacific Ocean from Peru to Alaska Bay (Kaas & Van Belle 1985 b; Sirenko et al. 2013). The fossil record includes Eocene of France, U. K., Ukraine and Hungary (Bielokrys 1999; Dell’Angelo et al. 2011, 2015 b; Dulai et al. 2017), upper Eocene or lower Oligocene of Washington, U. S. A. (Dell’Angelo et al. 2011), Oligocene of France, Germany, U. S. A. (Washington) and Canada (Janssen 1978; Dell’Angelo et al. 2019 b, 2020 b), and Miocene-Pleistocene of Europe and the Mediterranean Basin (Garilli et al. 2005; Studencka & Dulai 2010; Dell’Angelo et al. 2013, 2016, 2018 b). ...... continued on the next page	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF274EDB0FADFA126E7C94B4.taxon	discussion	Remarks. The genus Lepidochitona is well represented in the Paleogene and Neogene of Europe. The comparative analysis of these fossil species is usually hampered by the poor or incomplete descriptions available for many nominal taxa attributed to this genus; furthermore, a SEM-documentation of the ornamentation to unravel diagnostic details of their microsculpture is often lacking, and original descriptions are usually quite generic to allow a robust taxonomic identification. The genus Lepidochitona houses the greater number of species for any single genus treated in this study, 29 in all; to best facilitate their identification, we have grouped them by source area.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF244EDD0FADFD446B0995C4.taxon	description	Fig. 100	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF244EDD0FADFD446B0995C4.taxon	materials_examined	Type material. Lectotype MCZR E 20 / 12698 (Monterosato coll.), designated by Gaglini (1985), figured by Gaglini (1985: pl. 1, figs 1 – 2) and Cretella et al. (2005: fig. 1 a). Type locality. Capri Island (Campania, Italy). Material examined. Upper Miocene (Tortonian): Italy: Piedmont: Borelli: 2 valves (BD 837, MZB 32092, Fig. 100 D), Rio di Bocca d’Asino: 13 valves (BD 838, MZB 32090 - 32091, PG, Figs 100 E – F); Emilia-Romagna: Montegibbio: 2 valves (BD 839, LB). Lower Piocene: Italy: Liguria: Borzoli: 6 valves (BD 840, MZB 45755), Genova Sestri: 3 valves (BD 841), Rio Sant’ Antonino: 48 valves (BD 842, MP, MZB 45752 – 45754, Figs 100 A – B, 100 I), Zinola: 1 valve (BD 843). Pliocene: Italy: Tuscany: Orciano Pisano: 1 valve (BD 844), Poggio alla Fame: 5 valves (BD 845, Fig. 100 C), Serre di Rapolano: 11 valves (BD 846); Sicily: Altavilla: 2 valves (AG, AR), Trappeto: 1 valve (BD 847). Pleistocene: Italy: Tuscany: Riparbella: 15 valves (BD 848); Emilia-Romagna: Torrente Guerro: 1 valve (BD 849), Torrente Stirone: 8 valves (BD 850); Puglia: Gallipoli: 20 valves (BD 851); Calabria: Archi S. Francesco: 2 valves (BD 852, Figs 100 J – L), Le Castella: 3 valves (BD 853), Musalà: 1 valve (BD 854), Pecoraro: 3 valves (BD 855), Pezzo: 14 valves (BD 856); Sicily: Capo Milazzo: 1 valve (BD 857). Recent: Cyprus: Limassol: valves (BD 858, Figs 100 G – H). Maximum width of the valves: 2.2 / 3.4 / 1.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF244EDD0FADFD446B0995C4.taxon	description	Description. Head valve a little less than semicircular, with 8 – 10 radial depressions crossing its surface, front slope straight. Intermediate valves broadly rectangular (W / L = 2.57 – 2.90), moderately elevated (H / W = 0.35 – 0.48), rounded to semicarinate in anterior profile, anterior margin almost straight to slightly concave, side margins rounded, posterior margin almost straight, with a small apex in juvenile specimens, tending to disappear in adults, lateral areas hardly raised, though indicated by a rounded diagonal rib, another subsolete rib along the posterior margin. Tail valve triangular, W / L = 2.21, anterior margin almost straight to slightly sinuose, mucro subcentral, elevated, antemucronal slope almost straight, postmucronal slope almost straight to slightly concave. Tegmentum uniformly covered with rather coarse rounded and elevated granules (diameter ca 100 µm) arranged in irregular quincunxes. Each granule with one central megalaesthete surrounded by 15 – 20 micraesthetes. Articulamentum with apophyses wide, triangular in intermediate valves, rectangular in tail valve, teeth thickened at edges, those of tail valve very short, rugose, somewhat directed forward, slit formula 8 – 10 / 1 / 10 – 12, slits short, inequidistant, slit rays clearly visible, eaves coarsely spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF244EDD0FADFD446B0995C4.taxon	discussion	Remarks. The fossil record of Lepidochitona caprearum (Scacchi, 1836) is known from the upper Miocene of N. Italy, the Pliocene of Italy and Greece and the Pleistocene of Italy. This species is rather rare as a fossil, and has a long and rather complex nomenclatural history, summarized by Dell’Angelo & Smriglio (1999). Comparisons. Lepidochitona caprearum is close to L. canariensis (Thiele, 1909), for the differences see under this species (below).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF244EDD0FADFD446B0995C4.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Borelli, Montegibbio, Rio di Bocca d’Asino (Laghi 1977; Dell’Angelo et al. 1999, 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli. Genova Sestri, Rio S. Antonino, Zinola (Dell’Angelo et al. 2013; Dell’Angelo et al. 2021 b). Pliocene: central Mediterranean, Italy: Altavilla, Orciano Pisano, Poggio alla Fame, Serre di Rapolano, Trappeto (Laghi 1977; Dell’Angelo et al. 2001 a, 2012; Chirli 2004; this study). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece (Koskeridou et al. 2009). Pleistocene: central Mediterranean, Italy: many localities (Dell’Angelo et al. 2001 a, 2007 b; this study). Recent: Atlantic Ocean: along the coasts of France (Lacourt 1977), Spain and Portugal and in the Selvagens Arch. (Kaas 1991; Consolado Macedo et al. 1999; Carmona Zalvide et al. 2000); Mediterranean Sea: Alboran isl. (Salas & Luque 1986), France (B. D. D. 1882), Italy (Tiberi 1877; Dell’Angelo & Smriglio 1999; Trono 2006), Croatia (Dell’Angelo & Zavodnik 2004), Aegean Sea (Strack 1990; Koukouras & Karachle 2005), Tunisia (Kaas 1989; Cecalupo et al. 2008), Lebanon (Crocetta et al. 2014), Turkey (Ozturk et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF224EDF0FADFED868A19700.taxon	description	Fig. 101	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF224EDF0FADFED868A19700.taxon	materials_examined	Type material. LSL (fide Dodge 1952: 23). Type locality. “ In O. Norvegico ”. Material examined. Upper Miocene (Tortonian): Italy: Piedmont: Borelli: 2 valves (MZB 32089, MGPT, Fig. 101 B), Rio di Bocca d’Asino: 42 valves (BD 859, Figs 101 E – G, MZB 32086, Figs 101 J – L, PG), Villa Monti: 2 valves (BD 860); Emilia-Romagna: Montegibbio: 18 valves (BD 861, MZB 32087 – 32088, Fig. 101 A). Lower Pliocene: Italy: Liguria: Borzoli: 2 valves (BD 862, Figs 101 H – I, MZB 45750); Bussana: 2 valves (BD 863); Caranchi: 2 valves (BD 864, MZB 45751). Pliocene: Portugal: Vale de Freixo: 93 valves (BD 244, GeoFCUL VFX. 03.348, GeoFCUL VFX. 03.358 – VFX. 03.360, MNHN. F. A 81990, RGM. 1364016). Italy: Piedmont: Belveglio: 1 valve (BD 865); Tuscany: Bibbiano: 5 valves (BD 866), Montenero: 5 valves (BD 867), Orciano Pisano: 3 valves (BD 868), Pietrafitta: 11 valves (BD 869), Pietrafitta Melograni: 7 valves (BD 870), San Miniato: 5 valves (BD 871), Serre di Rapolano: 18 valves (BD 872); Emilia-Romagna: Castell’Arquato: 2 valves (BD 873), Cava di Campore: 1 valve (BD 874), Marano sul Panaro: 1 valve (BD 875); Sicily: Trappeto: 3 valves (BD 876). Pleistocene: Italy: Tuscany: Caletta: 1 valve (BD 877), Cisternino: 2 valves (BD 878), Riparbella: 7 valves (BD 879); Emilia-Romagna: Torrente Stirone: 5 valves (BD 880); Calabria: Archi S. Francesco: 6 valves (BD 881), Bovetto: 5 valves (BD 882), Carrabbati: 2 valves (BD 883), Musalà: 4 valves (BD 884), Pecoraro: 1 valve (BD 885), Pezzo: 5 valves (BD 886), Saracinello: 1 valve (BD 887), Stalettì: 2 valves (BD 888), Torrente Boscaino: 1 valve (BD 889); Sicily: Capo Milazzo: 2 valves (BD 890), Selinunte: 2 valves (BD 891). Greece: Kyllini: 3 valves (DGUP, Figs 101 C – D). Maximum width of the valves: 4.5 – 5 – 2.6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF224EDF0FADFED868A19700.taxon	description	Description. Head valve semicircular, posterior margin markedly V-shaped, front slope straight. Intermediate valves broadly rectangular (W / L = 2.42 – 2.77), moderately elevated (H / W = 0.30 – 0.45), semicarinate in anterior profile, anterior margin straight to slightly sinuose, side margins rounded, posterior margin almost straight, produced to a small apex, not beaked, lateral areas slightly raised, not distinctly separated from central area. Tail valve semicircular (W / L = 1.80 – 1.84), anterior margin slightly sinuose, mucro subcentral, not prominent, antemucronal slope straight, postmucronal slope slightly concave. Tegmentum uniformly covered with fine subquadrangular granules (maximum length 50 – 65 µm), arranged in somewhat irregular quincunx pattern, sometimes giving the impression of forming longitudinal striae towards the side margins, a few concentric growth lines often discernible. Each granule bearing a subcentral megalaesthete, several micraesthetes (up to 20) irregularly arranged along margin, also in depressions between granules. Articulamentum with apophyses wide, rounded, subtrapezoidal in tail valve, slit formula 8 – 10 / 1 / 8 – 12, slits inaequidistant, slit rays clearly visible, teeth short, slightly roughened on outside, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF224EDF0FADFED868A19700.taxon	discussion	Remarks. This species has a rather complicated taxonomic history, resulting in a great number of synonyms (Kaas & Van Belle 1981; Sirenko et al. 2016). Baxter (1982) studied the high variability of some characters of modern Atlantic individuals, such as number of incisions in the insertion laminae, ranging between 7 and 11 for the head valve and between 6 and 16 for the tail one. Baxter & Jones (1981) analysed the valve structure and the size and distribution of aesthetes Comparisons. The closest species is Lepidochitona pliocinerea Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013, and the differences with this species are discussed below.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF224EDF0FADFED868A19700.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian-Messinian): Po Basin, N Italy: Borelli, Montegibbio, Rio di Bocca d’Asino, Villa Monti (Sacco 1897; Laghi 1977; Dell’Angelo et al. 1999, 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Caranchi (Dell’Angelo et al. 2013, 2021 b). Pliocene: northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo & Silva 2003; Dell’Angelo et al. 2022); central Mediterranean, Italy: many localities in Piedmont, Tuscany, Emilia-Romagna, Sicily (Dell’Angelo et al. 2001 a; Chirli 2004; this study); Greece: Kallithea (Kleinhölter 1994). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece: Rhodes Island Koskeridou et al. 2009). Pleistocene: North Europe: Netherlands (Strack 2010); northeastern Atlantic: U. K. (Norton 1967), Portugal: Douro (Nobre 1892); central Mediterranean, Italy: many localities in Emilia-Romagna, Tuscany, Calabria, Sicily (Taviani 1978; Dell’Angelo et al. 2001 a, 2007 b; this study), Greece: Kyllini (Garilli et al. 2005). Holocene: North Atlantic, Wadden Sea, Denmark, Germany (Funder et al. 2002). Recent: Atlantic Ocean: from Norway: Tromsö (Sirenko 1998), Iceland (Sneli & Gudmundsson 2018), all the European coast (Leloup 1934; Rasmussen 1973; McKay & Smith 1979; Kaas & Van Belle 1981; Hansson 1998), France (Van Belle 1972); Spain and Portugal (Borja 1987; Consolado Macedo et al. 1999; Carmona Zalvide et al. 2000; Urgorri et al. 2017), Berlengas Arch. (Pisani Burnay 1986); Morocco, Senegal (Kaas 1991); Mediterranean Sea: Spain (Giribet & Penas 1997); France (B. D. D. 1882); Italy: many localities (Dell’Angelo & Smriglio 1999; Trono 2006); Malta (Mifsud et al. 1990); Greece and Aegean Sea Islands (Strack 1988, 1990; Koukouras & Karachle 2005); Turkey (Ozturk et al. 2014); Israel (Barash & Danin 1977); Algeria (Pallary 1900); Tunisia: Gulf of Gabes (Kaas 1989; Cecalupo et al. 2008); Black Sea and Sea of Marmara (Strack 1988; Bitlis 2019; Sirenko 2023).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF204EDF0FADFD1C6A9492DC.taxon	description	Fig. 102	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF204EDF0FADFD1C6A9492DC.taxon	description	(database: chiton fossil records); Koskeridou et al. 2009, p. 320, figs 10.6 – 10.7.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF204EDF0FADFD1C6A9492DC.taxon	materials_examined	Type material. Lectotype MCZR – M – 12718 / L, designated by Gaglini (1985). Paralectotype MCZR – M – 12718 / P (Appolloni et al. 2018). Type locality. Palermo (Sicily, Italy). Material examined. Pleistocene: Greece: Kyllini: 28 valves (BD 892, Figs 102 D – E, 102 G – H, DGUP, Figs 102 A – C, 102 F). Maximum width of the valves: - / 2.1 / 2.3 mm	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF204EDF0FADFD1C6A9492DC.taxon	description	Description. Head valve almost semicircular, posterior margin forms a very obtuse angle. Intermediate valves broadly rectangular (W / L = 2.73), rather depressed (H / W = 0.25 – 0.29), semicarinate in anterior profile, anterior margin straight, side margins rounded, posterior margin with pronounced apex, slightly concave at both sides, lateral areas hardly raised. Tail valve semielliptical (W / L = 1.70 – 2.01), anterior margin almost straight, mucro anterior, not prominent, antemucronal slope almost straight, postmucronal slope slightly concave. Tegmentum minutely granulose, almost smooth, with several concentric growth marks scarcely visible, aesthetes very dense. Articulamentum with apophyses triangular in intermediate valves, trapezoid in tail valve, separate by a jugal sinus flat, about one third of the valve’s width, teeth roughened at outside, slit formula 8 / 1 / 10 – 13, slits short, inequidistant, eaves very porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF204EDF0FADFD1C6A9492DC.taxon	discussion	Remarks. The fossil record of Lepidochitona furtiva (Monterosato, 1879) is limited to the Pleistocene of Greece (Garilli et al. 2005) and Rhodes (Koskeridou et al. 2009). Recently, several living specimens of Lepidochitona furtiva were found attached to thalli of Posidonia oceanica (Linnaeus) Delile, washed ashore following storms (Kaas & Van Belle 1988). This observation suggests an ecological association with the SGCF biocenosis (Gravina et al. 1992). A similar pattern is evident in the fossil record at Kyllini (Garilli et al. 2005), where numerous valves were recovered from the N 2 layer, which is notably rich in Posidonia remains, including both leaves and rhizomes. Comparisons. Lepidochitona furtiva shows some resemblance to L. verrucosa Dell’Angelo & Forli, 1996, a species known from the Tuscan Pliocene, that is characterized by a tegmentum covered with large irregular scales similar to warts (see below).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF204EDF0FADFD1C6A9492DC.taxon	distribution	Distribution. Upper Pliocene to upper Pleistocene: Greece: Rhodes Island (Koskeridou et al. 2009). Pleistocene: central Mediterranean, Greece: Kyllini (Garilli et al. 2005). Recent: Mediterranean: France, Italy (Dell’Angelo & Smriglio, 1999), Malta, Turkey (Ozturk et al. 2014) Algeria, Tunisia (Kaas 1989; Cecalupo et al. 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3F4EC10FADFCE76819915A.taxon	description	Fig. 103	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3F4EC10FADFCE76819915A.taxon	materials_examined	Type material. Holotype: MZB 11655, intermediate valve. Paratypes: MZB 11656 (3 valves); MZB 12755 – 12756 (2 valves); Museo di Storia Provinciale di Livorno (3 valves); MNHN (4 valves); private collections (19 valves): BD P 224 / 1 (6 valves, Figs 103 A – H), FG (7 valves), VB 3020 a (6 valves). Type locality. Capraia Island-Capo Corso, Ligurian Sea, from sediments at - 350 / 500 m trawled by fishers. Type stage. Pleistocene (“ biocenosi tardo quaternarie, in parte almeno dell’ultimo glaciale ”: late Quaternary biocoenoses, partly at least of last glacial age). Material examined. Pleistocene: Italy: Capraia Island, offshore of Capo Corso: type material, plus 259 valves (BD 893). Maximum width of the valves: 3.3 / 4.2 / 2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3F4EC10FADFCE76819915A.taxon	description	Description. Head valve a little less than semicircular, with 9 – 10 radial depressions crossing its surface, posterior margin forms a very obtuse angle, front slope slightly convex. Intermediate valves broadly rectangular (W / L = 2.40 – 2.79), moderately elevated (H / W = 0.32 – 0.48), semicarinate in anterior profile, anterior margin slightly concave, side margins rounded, posterior margin straight, with a protruding and well evident apex, lateral areas just raised, practically indistinguishable from central area. Tail valve less than semicircular (W / L = 2.07), anterior margin practically straight between apophyses, mucro subcentral, not prominent, antemucronal slope almost straight or slightly convex, postmucronal slope straight. Tegmentum rather coarse, uniformly covered with roundish and elevated granules (diameter ca 60 – 65 µm) arranged in irregular quincunxes; each granule with one central megalaesthete surrounded by up to 15 micraesthetes. Articulamentum with apophyses wide, more or less triangular, short and well incised teeth, much coarser, thicker and wrinkled in tail valve, slit formula 8 / 1 / 7 – 9, slits inequidistant, slit rays clearly visible, eaves coarsely spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3F4EC10FADFCE76819915A.taxon	discussion	Remarks. The material here discussed is the one described by Dell’Angelo & Giusti (1997), consisting of numerous loose valves trawled between 350 / 500 m offshore Capo Corso (Corsica) and Capraia Island, likely pertaining to glacial Pleistocene assemblages. Comparisons. The two closest species are Lepidochitona caprearum (Scacchi, 1836) and L. canariensis (Thiele, 1909), from which L. marcoi Dell’Angelo & Giusti, 1997 differs by the rather pronounced apex of the intermediate plates (not very evident in L. caprearum), LA practically indistinguishable from CA (with diagonal folds more or less evident in L. caprearum and L. canariensis), the more straight postmucronal slope of tail valve, the smaller granules and above all the different stratigraphic distribution, limited to the Pleistocene, presumably last glacial, for L. marcoi, broader, from Miocene to Recent, for L. caprearum and L. canariensis.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3F4EC10FADFCE76819915A.taxon	distribution	Distribution. Pleistocene: central Mediterranean, Italy: Capraia Island-Capo Corso, - 350 / 500 m (Dell’Angelo & Giusti 1997; Dell’Angelo et al. 2001 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3E4EC30FADFB2A6BF19698.taxon	description	Fig. 104	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3E4EC30FADFB2A6BF19698.taxon	materials_examined	Type material. Holotype: RMNH 55386 / 1; one specimen, length 7.8 mm. Paratypes: VB 2587 a / 1: one specimen disarticulated from Toulon, Le Carnier (France); VB 2587 b / 1: one specimen from Torba (Turkey); K 4849 / 1; one specimen from Torba (Turkey); MNHN / 1: one specimen from Torba (Turkey); IRSN 26095 / 1: one specimen from Tuscan Archipelago (Italy). Type locality. Cap d’Antibes, Port de l’Olivette (France), on a stone on sandy bottom, 0.5 m. Material examined. Upper Miocene: Italy: Montegibbio: 7 valves (BD 895, MZB 32096), Rio di Bocca d’Asino: 16 valves (BD 894, MZB 32094 – 32095, Figs 104 C – D, PG). Lower Pliocene: Italy: Liguria: Borzoli: 4 valves (BD); Bussana: 9 valves (BD 896); Caranchi: 1 valve (MZB 45758); Rio Sant’Antonino: 15 valves (BD 897, Figs 104 A – B, MP, MZB 45759 – 45760, Figs 104 K – L). Pliocene: Italy: Tuscany: Colle Val d’Elsa: 6 valves (BD 898), Montenero: 2 valves (BD 899), Orciano Pisano: 6 valves (BD 900), Pietrafitta Melograni: 1 valve (BD 901), Poggio alla Fame: 5 valves (BD 902, Figs 104 H – J). Pleistocene: Italy: Tuscany: Cisternino: 5 valves (BD 903), Riparbella: 5 valves (BD 904); Calabria: Le Castella: 3 valves (BD 905), Pecoraro: 2 valves (BD 906), Torrente Boscaino: 1 valve (BD 907); Sicily: Capo Milazzo: 2 valves (BD 908). Recent: Ustica Island: valves (BD 909, Figs 104 E – G). Maximum width of the valves: 2 / 2.5 / 2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3E4EC30FADFB2A6BF19698.taxon	description	Description. Head valve semicircular, with 8 radial depressions crossing its surface, posterior margin forms a very obtuse angle. Intermediate valves broadly rectangular (W / L = 2.04 – 2.27), moderately elevated (H / W = 0.53 – 0.57), semicarinate in anterior profile, anterior margin slightly concave between the apophyses, side margins rounded, posterior margin with strongly protruded apex, lateral areas conspicuously raised. Tail valve elliptical (W / L = 1.92), anterior margin slightly convex, mucro subcentral, not prominent, antemucronal slope almost straight, postmucronal slope slightly concave. Tegmentum rather coarsely granulated, granules roundish / oval (diameter up to 60 – 75 µm), convex, arranged in quincunx on HV, LA and PMA, and also on JA; arranged in curved and diverging longitudinal series on PA and AMA. Each granule with a central megalaesthete, and several micraesthetes (up to 15 – 20) irregularly arranged along margin, also in depressions between granules. Articulamentum with apophyses broadly triangular in intermediate valves, trapezoid in tail valve, separate by a jugal sinus about one third of the valve’s width, teeth obtuse, rugose on outside, slit formula 8 / 1 / 11, slits short, inequidistant, slit rays clearly visible, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3E4EC30FADFB2A6BF19698.taxon	discussion	Remarks. The species is rather rare as a fossil, with scarce records from the Italian Miocene, whilsts is commoner in the Plio-Pleistocene. Complete valves are rare, many of the examined valves are small fragments, in poor conditions of preservation, which makes specific attribution more difficult. Dell’Angelo et al. (2016) attributed tentatively to Lepidochitona monterosatoi Kaas & Van Belle, 1981 two incomplete valves (head and intermediate) from the Middle Miocene (Serravallian) of Monchio di Sarzano Casina (Italy). Despite the existing similarity with L. monterosatoi, it is not possible to identify with certainty these valves, that are discussed and figured in the section dedicated to the “ species of problematic assignment ”, and are left in open nomenclature as Lepidochitona sp. The original description of this species by Kaas & Van Belle reports H / W = 0.41, while the fossil material examined shows a higher value, from 0.53 to 0.57. On the other hand, also the living intermediate valves from Favignana Island (Dell’Angelo & Smriglio 1999: pl. 53, fig. I) and from Ustica Island (Figs 104 E – G) shows H / W values within the range of variation of fossil valves. The living specimens recorded from the Red Sea (Dahlak islands) by Leloup (1980), as Middendorffia caprearum (Scacchi, 1836), were later ascribed to L. monterosatoi by Strack (1993). The validity of this unique record outside the typical range needs to be reconsidered. Comparisons. Lepidochitona monterosatoi Kaas & Van Belle, 1981 is easily distinguishable from the other Mediterranean Lepidochitona by the intermediate valves highly elevated, the conspicuously raised lateral areas, and the arrangement of the granules on the pleural parts of the intermediate valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3E4EC30FADFB2A6BF19698.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Montegibbio, Rio di Bocca d’Asino (Dell’Angelo et al. 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: Borzoli, Bussana, Caranchi, Rio S. Antonino (Dell’Angelo et al. 2001 a, 2013, 2021 b). Pliocene: central Mediterranean, Italy: Tuscany: Colle Val d’Elsa, Montenero, Orciano Pisano, Pietrafitta Melograni, Poggio alla Fame (Forli et al. 1999; Dell’Angelo et al. 2001 a, 2013; Chirli 2004; this study). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece: Rhodes Island (Koskeridou et al. 2009). Pleistocene: central Mediterranean, Italy: Tuscany: Cisternino, Riparbella (Dell’Angelo & Forli 1995 a; this study), Calabria: Le Castella, Pecoraro, Torrente Boscaino (this study), Sicily: Capo Milazzo (this study). Recent: Atlantic Ocean: Spain and Portugal (Carmona Zalvide et al. 2000; Urgorri et al. 2017). Mediterranean Sea (Dell’Angelo & Smriglio 1999; Koukouras & Karachle 2005; Cecalupo et al. 2008; Crocetta et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3C4EC40FADFC64684E92A3.taxon	description	Fig. 105	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3C4EC40FADFC64684E92A3.taxon	materials_examined	Type material. Holotype: MZB 49979, intermediate valve, Figs 105 F – G. Paratypes: MZB 49977 (1 valve from Bussana); MZB 49978 (1 valve from Rio Sant’Antonino, Figs 105 A – B); MSNG 56537 (2 valves from Bussana and Rio Sant’Antonino); private collections (4 valves): BD: (2 valves from Serre di Rapolano, Fig. 105 H), MP (1 valve from Rio S. Antonino), MS (1 valve from from Genova Sestri). Type locality. Bussana (Liguria, Italy). Type stage. Lower Pliocene (Zanclean). Material examined. Lower Miocene: Italy (Burdigalian): Piedmont: Sciolze: 1 valve (PG). Middle Miocene: Italy (Langhian): Piedmont: Albugnano: 1 valve (MZB 32097, Figs 105 C – D). Upper Miocene: Italy (Tortonian): Piedmont: Borelli: 5 valves (BD 910, MGPT PU 135299, Fig. 105 E), Rio di Bocca d’Asino: 21 valves (BD 911, MZB 32098, Fig. 105 L, PG); Emilia-Romagna: Montegibbio: 7 valves (BD 912, MZB 32099). Lower Pliocene: Italy: Liguria: type material, plus Borzoli: 14 valves (BD 913); Bussana: 10 valves (BD 914); Caranchi: 4 valves (BD 915, MP); Garlenda: 2 valves (MP); Genova Sestri: 2 valves (BD 916); Rio Sant’Antonino: 75 valves (BD 917, MP). Pliocene: Italy: Tuscany: Serre di Rapolano (21 valves) (BD 918, Figs 105 I – K); Spain: Estepona: 6 valves (BD 919). Maximum width of the valves: 7.5 – 15 – 6.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3C4EC40FADFC64684E92A3.taxon	description	Description. Valves solid, often remarkably thick. Head valve about one third of a circle. Intermediate valves broadly rectangular (W / L = 2.83 – 3.74), moderately elevated (H / W = 0.31 – 0.35), semicarinate in anterior profile, posterior margin almost straight, drawn out to small apex, not beaked, lateral areas not distinctly separated from central area. Tail valve elliptical (W / L = 2.04 – 2.35), anterior margin almost straight or slightly convex, mucro subcentral, not prominent, antemucronal and postmucronal slopes practically straight. Tegmentum evenly sculptured all over with irregularly roundish granules (diameter up to 115 µm), arranged in somewhat irregular quincunx pattern. Each granule bearing more or less central megalaesthete and several micraesthetes (15 – 20) irregularly arranged along margin, also in depressions between granules. Articulamentum with apophyses short, very wide, starting from limits of jugal area, jugal sinus straight to slightly convex, narrow about one seventh of total width. Slit formula 15 – 16 / 1 / 11 – 13, slit rays well marked, insertion teeth inaequidistant.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3C4EC40FADFC64684E92A3.taxon	discussion	Remarks. Most material is represented by small fragments, whilst complete valves are rare. The material reported as Lepidochitona cinerea (Linnaeus, 1767) from the Pliocene of Estepona (Dell’Angelo et al. 2004), and some of the valves from the Pliocene of Tuscany (Chirli 2004, partim; Dell’Angelo et al. 2001 a, partim) has been ascribed to L. pliocinerea by Dell’Angelo et al. (2013). Comparisons. Lepidochitona pliocinerea Dell’Angelo, Sosso, Prudenza & Bonfitto, 2013 is very similar to L. cinerea (Linnaeus, 1767), from which it differs by the greater width and the more elongate shape of its valves, the larger apophyses starting from the limits of JA, the different slope in tail valve, the greater number of slits (slit formula 15 – 16 / 1 / 11 – 13 vs. 8 – 10 / 1 / 8 – 12 in L. cinerea), and the more irregularly roundish granules (vs. subquadrangular and more regular in L. cinerea).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3C4EC40FADFC64684E92A3.taxon	distribution	Distribution. Lower Miocene: Proto-Mediterranean Sea (Burdigalian): N. Italy: Sciolze (Dell’Angelo et al. 2016). Middle Miocene: Proto-Mediterranean Sea (Langhian): N. Italy: Albugnano (Dell’Angelo et al. 2016). Late Miocene: Proto-Mediterranean Sea (Tortonian): N. Italy: Borelli, Montegibbio, Rio di Bocca d’Asino (Dell’Angelo et al. 2016). Lower Pliocene: central Mediterranean, Italy: Liguria (Dell’Angelo et al. 2013, 2021 b). Pliocene: western Mediterranean: Spain: Estepona (Dell’Angelo et al. 2004), central Mediterranean: Italy: Tuscany: Serre di Rapolano (Dell’Angelo et al. 2001 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	description	Fig. 106	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	materials_examined	Type material. Holotype: MSNG 62664, intermediate valve, width 4.5 mm (Figs 106 A – F). Type locality. Pezzo, near Villa S. Giovanni (Calabria, Italy). Type stage. Upper Pleistocene, possibly originated from contiguous Lower Pleistocene bathyal deposits.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	etymology	Etymology. The name honours Hiroshi Saito (National Museum of Nature and Science, Japan), for his prominent contribution to the study of Recent and fossil chitons.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	materials_examined	Material examined. Pezzo: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	diagnosis	Diagnosis. Intermediate valve solid, triangular, strongly beaked, elevated, semicarinate in anterior profile, posterior margin straight on both sides of the prominent apex, lateral areas strongly raised. Tegmentum coarsely granulated, fully covered by roundish / oval granules. Articulamentum with apophyses broadly triangular, slit formula one slit for each side.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	description	Description. Head and tail valves unknown. Intermediate valve solid, triangular (W / L = 1.44), strongly beaked, elevated (H / W = 0.51), semicarinate in anterior profile, anterior margin convex, posterior margin straight on both sides of the prominent apex, forming an angle of 120 °, lateral areas strongly raised, delimited from central area by a deep depression. Tegmentum coarsely granulated, fully covered by granules roundish / oval, irregularly arranged, LA with strong, concentric folds. Articulamentum with apophyses broadly triangular, separate by a large jugal sinus, slit formula one slit for each side, slit rays visible, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	discussion	Remarks. The species is known only from the single intermediate valve from Pezzo (Italy) here described, with a characteristic shape. Comparisons. This species shows a valve’s shape closest to that of Lepidochitona modesta (Rolle, 1862), a species from the Oligocene of Gaas (France), also known for a single intermediate valve (see below), but with a different sculpture (smooth).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC50FADFF056EF793C2.taxon	distribution	Distribution. Upper Pleistocene: central Mediterranean, S. Italy: Pezzo (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC60FADF8D16BD79394.taxon	description	Fig. 107	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC60FADF8D16BD79394.taxon	materials_examined	Type material. Holotype: MZB 11653, 1 intermediate valve. Paratypes: MZB 12696 – 12697 (2 valves, fide Dell’Angelo et al. 2001 a); DSTUT B-P. I. 06.1 (3 valves); private collections (9 valves): BD F 55 D / 1 (3 valves, 2 from Pietrafitta Melograni and 1 from Pietrafitta “ Sbarra ”), MF (3 valves), EU (3 valves). Type locality. Pietrafitta Melograni (Tuscany, Italy). Type stage. Pliocene. Material examined. Pliocene: Italy: Tuscany: type material, plus Aiano: 1 valve (BD 920), Bibbiano: 4 valves (BD 921), Colle Val d’Elsa: 5 valves (BD 922), Orciano Pisano: 4 valves (BD 923), Pietrafitta: 62 valves (BD 924), Pietrafitta Melograni: 84 valves (BD 925, Figs 107 B – D), Poggio alla Fame: 10 valves (BD 926, Figs 107 A, 107 E – H). Maximum width of the valves: 1.7 – 2.2 – 1.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC60FADF8D16BD79394.taxon	description	Description. Head valve almost semicircular. Intermediate valves broadly rectangular (W / L = 2.38 – 2.50), rather depressed (H / W = 0.22 – 0.36), semicarinate in anterior profile, anterior margin straight, side margins rounded, posterior margin with pronounced apex, lateral areas hardly raised, almost indistinguishable from central area. Tail valve semielliptical, width twice length (W / L = 2.06), anterior margin straight, mucro subcentral, not prominent, antemucronal and postmucronal slopes almost straight. Tegmentum covered with large, irregular, wart-like scales, obliquely oriented and more or less uniformly distributed across the entire surface of the valves. The lateral areas often exhibit several well-marked growth striae. Articulamentum with apophyses triangular in intermediate valves, trapezoid in tail valve, teeth short, rough and well cut, of irregular width, at times with extra incisions seeming to further split real teeth, but stop half-way their length, slit formula 9 – 10 / 1 / 9 – 10, slits inequidistant, eaves very porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC60FADF8D16BD79394.taxon	discussion	Remarks. The fossil record of Lepidochitona verrucosa Dell’Angelo & Forli, 1996 is limited to a few localities from the Pliocene of Tuscany. Comparisons. This species is easily recognized for the tegmentum covered with large irregular scales similar to warts.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF3A4EC60FADF8D16BD79394.taxon	distribution	Distribution. Pliocene: central Mediterranean, Italy: Tuscany: Aiano, Bibbiano, Colle Val d’Elsa, Orciano Pisano, Pietrafitta, Pietrafitta Melograni, Poggio alla Fame (Dell’Angelo & Forli 1996; Dell’Angelo et al. 2001 a; Chirli 2004; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF394ECA0FADF97F6BFB95EE.taxon	description	personal observations).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF344ECB0FADFF056BBD92E7.taxon	description	Fig. 108	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF344ECB0FADFF056BBD92E7.taxon	materials_examined	Type material. Holotype: MHNBx 2020.5.1, tail valve, width 2 mm, Figs 108 F – H. Paratypes: MHNBx 2020.5.2, head valve, width 1.8 mm, Figs 108 A – B; MHNBx 2020.5.3, intermediate valve, width 2.4 mm, Figs 108 C – E. Type locality. Lariey (France). Type stage. Miocene (Aquitanian). Material examined. Lower Miocene (Aquitanian): Lariey: type material plus 12 valves (JFL); Pouquet: 1 head valve (JFL). Maximum width of the valves: 1.8 / 2.7 / 2.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF344ECB0FADFF056BBD92E7.taxon	description	Description. Head valve semicircular, apex recurved. Intermediate valves broadly rectangular, width twice length (W / L = 2.00 – 2.25), rounded in anterior profile, moderately elevated (H / W = 0.30 – 0.35), anterior margin slightly convex, lateral margins rounded, posterior margin concave at both sides of prominent apex, lateral areas scarcely differentiated. Tail valve semicircular (W / L = 1.64), weakly elevated, anterior margin slightly convex, mucro subcentral, antemucronal slope almost straight, postmucronal slope concave. Tegmentum smooth, displaying very fine microgranulation, aesthetes of same size. Articulamentum with apophyses small and trapezoidal, extending to lateral margins in intermediate valves, slit formula 8 / 1 / 11 – 12, teeth inequidistant.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF344ECB0FADFF056BBD92E7.taxon	discussion	Remarks. Lepidochitona adunca (Dell’Angelo, Lesport, Cluzaud & Sosso, 2020) is know only for two sites in the Aquitanian Basin (Pouquet, and Saucats) from the Miocene (Aquitanian). This species was initially attributed to the genus Tonicella Carpenter, 1873, and later included by Sirenko & Dell’Angelo (2023) in the genus Lepidochitona Gray, 1821, which, since the Eocene, was widespread in the Tethys Ocean. Apparently, Tonicella first appeared in the Atlantic Ocean during the Pliocene (Sirenko & Dell’Angelo 2023). Comparisons. Lepidochitona adunca differs from L. lira Cherns & Schwabe, 2019, mainly by the smooth surface (vs. lateral areas rugose in L. lira), posterior margin tapering to a pointed apex (vs. almost straight with apex not evident in L. lira), recurved apex in the head valve, and trapezoidal apophyses extending to lateral margins in intermediate valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF344ECB0FADFF056BBD92E7.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Aquitanian): Aquitaine Basin, France: Lariey, Pouquet (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF334ECD0FADFF05695E912C.taxon	description	Fig. 109	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF334ECD0FADFF05695E912C.taxon	materials_examined	Type material. Holotype ZMHU 101918. Type locality. Canary Islands, Tenerifa, Puerto. Material examined. Upper Miocene: France (Messinian?): Moulin-Pochas: 1 valve (PR, Fig. 109 I). Lower Pliocene: Italy: Liguria: Borzoli: 3 valves (BD 927, MZB 45757), Bussana: 1 valve (BD 928), Genova Sestri: 2 valves (BD 929, Figs 109 J – L), Rio Sant’ Antonino: 2 valves (MP, MZB 45756), Rio Torsero: 1 valve (BD 930, Figs 109 C – D). Pliocen e: Portugal: Vale de Freixo: 2 valves (lost during photography). Spain: Estepona: 2 valves (BD 931); Italy: Piedmont: Vintebbio: 5 valves (BD 932, Figs 109 E – H); Tuscany: Poggio alla Fame: 10 valves (BD 933, Figs 109 A – B). Pleistocene: Italy: Calabria: Gallina: 8 valves (BD 934). Maximum width of the valves: 2.5 / 4.8 / 1.4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF334ECD0FADFF05695E912C.taxon	description	Description. Head valve about three fifth of a circle, front slope straight. Intermediate valves broadly rectangular (W / L = 2.58 – 2.86), moderately elevated (H / W = 0.28 – 0.42), semicarinate in anterior profile, anterior margin concave between the apophyses, side margins rounded, posterior margin almost straight, with small but well pronounced apex, lateral areas hardly raised, though sometimes indicated by a faint diagonal fold. Tail valve about one third of a circle (W / L = 2.04 – 2.38), anterior margin almost straight, mucro subcentral or in anterior position, prominent, antemucronal slope almost straight, postmucronal slope slightly concave just under mucro. Tegmentum uniformly covered with diamond-shaped granules (maximum length 60 – 80 µm), arranged in irregular quincunxes, tending to form converging longitudinal rows on PA. Each granule with one central megalaesthete surrounded by 10 – 15 micraesthetes. Articulamentum with apophyses small, more or less triangular, teeth blunt, roughened on outside, slit formula 7 – 10 / 1 / 8 – 10, slits short, slit rays clearly visible, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF334ECD0FADFF05695E912C.taxon	discussion	Remarks. The species is rather rare as a fossil, scarce reports from the French Miocene of Ligerian Basin, more common from Pliocene and rare from the Italian Pleistocene. Dell’Angelo et al. (2020 b) attributed tentatively to Lepidochitona canariensis (Thiele, 1909) a single intermediate valve (Fig. 154 O – P) from the lower Miocene (Burdigalian) of the Aquitaine Basin. Despite a morphological similarity with L. canariensis, this identification is doubtful and is therefore left in open nomenclature as Lepidochitona sp. (see below, “ species of problematic assignment ”). Comparisons. Lepidochitona canariensis is closest to L. caprearum (Scacchi, 1836), from which it differs by the head valve without radial depressions (present in L. caprearum), the different shape of the tail valve, the sculpture with more irregular and smaller granules (more roundish in L. caprearum).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF334ECD0FADFF05695E912C.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic: Ligerian Basin, France: Moulin Pochas (Dell’Angelo et al. 2020 b). Lower Pliocene: central Mediterranean, Italy: Liguria: Bussana, Rio S. Antonino, Rio Torsero, Sestri Ponente, Borzoli (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013). Pliocene: northeastern Atlantic, Mondego Basin, Portugal (Dell’Angelo & Silva 2022). Western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: Piedmont: Vintebbio, Tuscany: Poggio alla Fame (this study). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece: Rhodes Island (Koskeridou et al. 2009). Pleistocene: central Mediterranean, S. Italy: Gallina (this study). Recent: Atlantic Ocean: Portugal, Spain (Carmona Zalvide et al. 2000); Mauritania: Agadir Island (Anseeuw & Verstraeten 2009), Madeira (Van Belle 1985; Kaas 1991), Selvagens Arch. (Segers et al. 2009), Cape Verde Arch. (Strack 2005), Azores (Avila & Sigwart 2013) and the Canary Islands (Leloup 1968; Van Belle 1984; Kaas 1991; Hernández & Rolán 2011). Mediterranean Sea: Morocco: Torres de Alcalà (Dell’Angelo & Tringali 2000).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF324ECE0FADFBF06B599661.taxon	description	Fig. 110	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF324ECE0FADFBF06B599661.taxon	materials_examined	Type material. Holotype: MHNBx 2020.2.7, intermediate valve, width 2.2 mm (Figs 110 A – D). Paratype: MHNBx 2014.14.22, Benoist collection, labeled Tonicia gaasensis de Rochebrune, 1882, intermediate valve, width 2.8 mm. Type locality. Gaas (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas: type material plus Rupelian levels: 1 intermediate valve (BD 935); Espibos: 1 intermediate valve (DA); Larrat: 59 intermediate valves (AC, DA, JCV). Maximum width: 3.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF324ECE0FADFBF06B599661.taxon	description	Description. Head and tail valves not present. Intermediate valve broadly rectangular (W / L = 1.57 – 1.82), carinate in anterior profile, highly elevated (H / W = 0.43 – 0.58), anterior margin convex, lateral margins rounded, posterior margin concave at both sides of very prominent apex, lateral areas scarcely differentiated. Tegmentum entirely covered by irregularly arranged, elevated, roundish granules, some coalescent, diameter up to 93 μm; each granule bearing a subcentral megalaesthete, and up to six micraesthetes arranged irregularly along margin, additional micraesthetes randomly placed between granules. Articulamentum with apophyses small, triangular, slit formula one slit for each side.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF324ECE0FADFBF06B599661.taxon	discussion	Remarks. Lepidochitona chalossensis Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 is characterized by the tegmentum entirely covered by irregularly arranged roundish granules, a carinate anterior profile of the intermediate valves, and very prominent apex. The valve from the Benoist collection at MHNBx (MHNBx 2014.14.22, see Dell’Angelo et al. 2020 b), labeled Tonicia gaasensis de Rochebrune, 1882, has been considered conspecific to Lepidochitona chalossensis by Dell’Angelo et al. (2020 b). Comparisons. The closest species is Lepidochitona larratensis Dell’Angelo, Lesport, Cluzaud & Sosso, 2020, from which L. chalossensis differs by the different shape of the intermediate valves (H / W = 0.43 – 0.58 vs. 0.56 – 0.62 in L. larratensis), the carinate anterior profile (vs. semicarinate in L. larratensis), and the convex anterior margin (vs. slightly sinuose in L. larratensis).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF324ECE0FADFBF06B599661.taxon	distribution	Distribution. Middle Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF314ECF0FADFC3C6B599420.taxon	description	Fig. 111	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF314ECF0FADFC3C6B599420.taxon	materials_examined	Type material. Holotype: MHNBx 2020.3.6, intermediate valve, width 3.8 mm, Figs 111 A – E. Paratypes: MHNBx 2014.14.23.5, intermediate valve present in the lot deposited in the Benoist collection, labeled Lepidopleurus daubrei; MZB 50571, topotypic intermediate valve, Figs 111 F – H. Type locality. Gaas, Larrat (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas: type material plus Rupelian levels: 1 intermediate valve, width 3.2 mm (BD 936); Larrat: 8 intermediate valves, maximum width 4 mm (AC).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF314ECF0FADFC3C6B599420.taxon	description	Description. Head and tail valves not present. Intermediate valve broadly rectangular (W / L = 1.56 – 1.78), rounded in anterior profile, highly elevated (H / W = 0.55 – 0.70), anterior margin almost straight, lateral margins slightly rounded, posterior margin slightly concave at both sides of prominent apex, lateral areas scarcely differentiated. Tegmentum entirely covered by irregularly arranged, weak elevated, triangular to oval granules, not coalescent, diameter up to 110 μm; each granule with a more or less central megalaesthete, and up to 6 micraesthetes arranged irregularly along margin, additional micraaesthetes present between granules. Articulamentum with small, rounded apophyses, slit formula two slits on each side, slit rays visible, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF314ECF0FADFC3C6B599420.taxon	discussion	Remarks. The valve from the Benoist collection at MHNBx (MHNBx 2014.14.23.5), labeled Lepidopleurus daubrei de Rochebrune, 1882, has been considered conspecific to Lepidochitona charlesi Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 by Dell’Angelo et al. (2020 b). The shell characters in this species shows little variability. Only one valve (Figs 111 F – H) is much higher (H / W = 0.70, while all the other valves examined have H / W = 0.55 – 0.61) and has a more rounded to semicarinate anterior profile. The second slit in the articulamentum is smaller and near the posterior margin, not clearly visible in the holotype, but well evident in the rest of the material examined (Fig. 111 G). Comparisons. Lepidochitona charlesi Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 is characterized by having the tegmentum entirely covered by irregularly arranged triangular to oval granules (Fig. 111 B), larger than the other species of Lepidochitona from Gaas [except L. tarbelliana Dell’Angelo, Lesport, Cluzaud & Sosso, 2020]. The closest species is Lepidochitona tarbelliana, from which L. charlesi differs by the different shape of the valves (W / L = 1.56 – 1.78 vs. 1.82 – 2.26 in L. tarbelliana), higher elevation (H / W = 0.55 – 0.70 vs. 0.39 – 0.47 in L. tarbelliana, and the triangular to oval granules, not coalescent.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF314ECF0FADFC3C6B599420.taxon	distribution	Distribution. Middle Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF304EF00FADFBBC6B59910A.taxon	description	Fig. 112	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF304EF00FADFBBC6B59910A.taxon	materials_examined	Type material. Holotype MHNBx 2020.2.6, intermediate valve, width 2.7 mm (Figs 112 A – D). Paratype MHNBx 2020.2.9, intermediate valve (Fig. 112 E). Type locality. Gaas, Larrat (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas: type material plus Benoist collection (MHNBx 2014.14.21.1, MHNBx 2014.14.21.2, Figs 112 F – H, labeled Tonicia waltebledi de Rochebrune, 1882); Rupelian levels: 3 intermediate valves (BD 937); Larrat: 60 intermediate valves (AC, DA, JCV). Maximum width: 4.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF304EF00FADFBBC6B59910A.taxon	description	Description. Head and tail valves not present. Intermediate valve broadly rectangular (W / L = 1.44 – 2.13), semicarinate in anterior profile, highly elevated (H / W = 0.56 – 0.62), anterior margin slightly sinuose, weakly concave in jugum, lateral margins rounded, posterior margin concave at both sides of very prominent apex, lateral areas scarcely differentiated. Tegmentum entirely covered by irregularly arranged, elevated, roundish granules, some coalescent, diameter up to 86 μm; each granule bearing a more or less central megalaesthete, and up to eight micraesthetes arranged irregularly along margin, additional micraesthetes randomly placed between granules. Articulamentum with apophyses small and triangular, slit formula one slit for each side, slit rays clearly visible, second slit ray lies close to posterior margin.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF304EF00FADFBBC6B59910A.taxon	discussion	Remarks. This species is characterized by the tegmentum entirely covered by irregularly arranged roundish granules, and the intermediate valves elevated, with a very prominent apex. The two valves from the Benoist collection at MHNBx (MHNBx 2014.14.21.1, MHNBx 2014.14.21.2), labeled Tonicia waltebledi de Rochebrune, 1882, have been considered conspecific to Lepidochitona larratensis Dell’Angelo, Lesport, Cluzaud & Sosso by Dell’Angelo et al. (2020 b). These valves show a slightly different shape, with higher values of W / L (2.13 – 2.20 vs. 1.94 of the type material) and similar H / W (0.58 – 0.61 vs. 0.56 – 0.62 of the type material). Comparisons. The closest species is Lepidochitona chalossensis Dell’Angelo, Lesport, Cluzaud & Sosso, 2020, see above.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF304EF00FADFBBC6B59910A.taxon	distribution	Distribution. Middle Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0F4EF10FADFB1A6F62904A.taxon	description	Fig. 113	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0F4EF10FADFB1A6F62904A.taxon	materials_examined	Type material. Unknown, the status of the collection Bial de Bellerade is currently unknown, with some material (no “ type ” specimens), recorded in the University of Talence (Bordeaux). Type locality. Le Thil (France). Type stage. Lower Miocene. Material examined. Lower Miocene: France (Aquitanian): Lariey: 2 valves (JFL, Figs 113 A – C). France (Burdigalian): Gamachot: 6 valves (BD 938, MZB 50573 - 50575, Figs 113 D – L). Maximum width of the valves: 2.6 – 3.5 – 1.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0F4EF10FADFB1A6F62904A.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, slope slightly convex. Intermediate valves broadly rectangular (W / L = 2.06 – 2.53), semicarinate in anterior profile, moderately elevated (H / W = 0.44 – 0.45), anterior margin slightly convex or weakly sinuose, side margins poorly rounded, posterior margin straight at both sides of prominent apex, lateral areas scarcely differentiated. Tail valve semicircular (W / L = 2.46), anterior margin almost straight, mucro in anterior position, antemucronal and postmucronal slopes almost straight. Tegmentum entirely covered by randomly arranged elliptical granules, although weak impression of order in radial striae on LA, radiating from area just next to apex. Articulamentum with large apophyses, triangular in intermediate valves, trapezoidal in tail valve, slit formula: 9 / 2 / 7, slits inequidistant, slit rays clearly visible in head and tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0F4EF10FADFB1A6F62904A.taxon	discussion	Remarks. The fossil record of Lepidochitona leognanensis (Cossmann & Peyrot, 1917) is limited to the lower Miocene of the Aquitaine Basin. This species is poorly known, scarcely recorded in the literature since its original description; one record from Rennes Dolfuss (1920) recorded and the material recently reported from Lariey by Cahuzac et al, 2012 (the two intermediate valves here figured, Figs 113 A – C). Comparisons. See Tab. 18, 19 for more comparison with the other Lepidochitona spp. from the NE Atlantic considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0F4EF10FADFB1A6F62904A.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Aquitanian-Burdigalian): France, Aquitaine Basin: Gamachot, Lariey, Le Thil (Cossmann & Peyrot 1917; Cahuzac et al. 2012; Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0E4EF20FADFA5B6A94915A.taxon	description	Fig. 114	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0E4EF20FADFA5B6A94915A.taxon	materials_examined	Type material. Holotype: ZSM Mol 20071428 C, intermediate valve, Figs 114 C – D; paratype: ZSM Mol 20071428 B, head valve, Figs 114 A – B. Type locality. Abbesse (France). Type stage. Upper Oligocene (Chattian). Material examined. No material, only descriptions and illustrations from the literature. Maximum width of the valves: 2.6 / 5.6 / -- mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0E4EF20FADFA5B6A94915A.taxon	description	Description. Tail valve unknown. Head valve semi-circular, moderately elevated, subcarinate, posterior margin nearly straight. Intermediate valves broadly rectangular (W / L = 2.45), moderately elevated (H / W = 0.38), semicarinate in anterior profile, anterior margin tapering outwards from slightly to moderately convex jugal area, side margins fairly straight, posterior margin nearly straight, apex not evident, lateral area slightly elevated, not greatly distinct from central area, with shallow longitudinal ridges and furrows, depressed above diagonal slit ray. Tegmentum smooth displaying very fine microgranulation, smooth rounded broad CA showing a slight narrow jugal elevation and straight side slopes, indistinct very low ridges or growth lines parallel to anterior margin rounding at fairly square corners into ridges and furrows of LA, aesthetes of same size. Articulamentum with triangular, rounded, long and wide apophyses, narrow jugal sinus, insertion plate rounded, short, slit formula 8 / 1 / --, slit rays visible, thin, spongy eaves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0E4EF20FADFA5B6A94915A.taxon	discussion	Remarks. Lepidochitona lira (Cherns & Schwabe, 2019) is known only for the original finding of 12 valves (2 head and 10 intermediate, ZSM Mol 20071428) from the upper Oligocene (Chattian) of Abbesse (France). This species was initially attributed to the genus Tonicella Carpenter, 1873 by Cherns & Schwabe (2019), and later included by Sirenko & Dell’Angelo (2023) in the genus Lepidochitona Gray, 1821. Comparisons. See Tab. 18, 19 for comparison with the other Lepidochitona spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0E4EF20FADFA5B6A94915A.taxon	distribution	Distribution. Upper Oligocene: NE Atlantic (Chattian): Aquitaine Basin, France: Abbesse (Cherns & Schwabe 2019).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0D4EF30FADFB39697E90C1.taxon	description	Fig. 115	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0D4EF30FADFB39697E90C1.taxon	description	p. 31, fig. 20; Sirenko & Dell’Angelo 2023, p. 278. Lepidochitona modesta; Sirenko & Dell’Angelo 2023, p. 280.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0D4EF30FADFB39697E90C1.taxon	materials_examined	Type material. Holotype NHMW 2011 / 0009 / 0001: 1 intermediate valve, width 2.4 mm (Figs 115 A – C). Type locality. Gaas (France). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene (Rupelian): Gaas: type material plus 1 head valve (NHMW 2011 / 0009 / 0005, Fig. 115 D), 2 intermediate valves (BD 939, MZB 50576, Fig. 115 H); Larrat: 16 intermediate valves (AC, DA, JCV, MHNBx 2020.3.8, Figs 115 E – G). Maximum width of the valves: 2.1 / 3.4 / -- mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0D4EF30FADFB39697E90C1.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valve triangular (W / L = 1.55 – 1.72, 1.55 in holotype), strongly beaked, semicarinate in anterior profile, highly elevated (H / W = 0.51 – 0.55, 0.55 in holotype), anterior margin convex, posterior margin almost straight on both sides of prominent apex, forming an angle of 101 – 106 ° (103 ° in holotype), lateral areas strongly delimited from central area by a diagonal fold. Tegmentum smooth, displaying very fine microgranulation, some growth lines often present, aesthetes of same size. Articulamentum with apophyses small and triangular, slit formula: 10 / 1 / --, with another slit just visible in intermediate valve, close to posterior margin, slit rays clearly visible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0D4EF30FADFB39697E90C1.taxon	discussion	Remarks. Lepidochitona modesta (Rolle, 1862) is a species described on the basis of a single intermediate valve, with a characteristic shape (Rolle, 1862: 215 “ beak - like, backwards highly elongated shape ”, translation by A. Kroh). This valve, as already noted by Rolle, could be considered as the valve ii (the first intermediate one), in several chiton species characterized by a different shape, more lengthened and beaked, and often for this reason described or illustrated separately from the other intermediate valves (e. g., Kaas et al. 2006: figs 147 – 150). This species was attributed to the genus Tonicella Carpenter, 1873 by Dell’Angelo et al. (2018 b), and later included by Sirenko & Dell’Angelo (2023) in the genus Lepidochitona Gray, 1821. The head valve preserved in NHMW (Fig. 115 D) has been considered by Dell’Angelo et al. (2020 b) conspecific to Lepidochitona modesta, due to its tegmental surface, smooth and with arrangement of aesthetes like that of the holotype (compare Dell’Angelo et al. 2020 b: figs 6 C and 6 G). Comparisons. The shape of the intermediate valves of Lepidochitona modesta is very characteristic and differs from the shape of all other Lepidochitona species discussed in this study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0D4EF30FADFB39697E90C1.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Rolle 1862; de Rochebrune 1882; Dell’Angelo et al. 2019 b, 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0C4EF40FADF9DF6B59921D.taxon	description	Fig. 116	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0C4EF40FADF9DF6B59921D.taxon	materials_examined	Type material. Holotype: MHNBx 2020.3.9, tail valve, width 2.3 mm (Figs 116 F – H). Paratypes: MHNBx 2020.1.3, head valve, width 3 mm (Figs 116 A – B); MHNBx 2020.1.4, intermediate valve, width 3.8 mm (Figs 116 C – E). Type locality. Gaas, Larrat (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas: type material plus 9 valves (BD 940); Lagouarde: 3 valves (DA); Larrat: 214 valves (AC, DA, JCV). Maximum width of the valves: 3 / 4.4 / 2.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0C4EF40FADF9DF6B59921D.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, front slope straight, slight presence of radial folds. Intermediate valves broadly rectangular, width more than twice the length (W / L = 2.05 – 2.50), semicarinate in anterior profile, moderately to highly elevated (H / W = 0.41 – 0.51), anterior margin slightly convex, almost straight in jugum, lateral margins rounded, posterior margin concave at both sides of prominent apex, lateral areas delimited from central area by diagonal fold, in some valves scarcely visible, second weaker diagonal fold present on posterior margin. Tail valve semicircular, width about twice the length (W / L = 1.93 – 2.10), anterior margin slightly convex, straight or slightly concave in jugal portion, mucro subcentral, antemucronal slope almost straight, postmucronal slope straight or slightly convex. Tegmentum smooth, glossy, with several inconspicuous growth lines, one or two more evident near anterior margin on head valve, aesthetes of same size. Articulamentum with apophyses triangular, trapezoidal in tail valve, weakly projecting, slit formula 8 – 9 / 1 / 8, insertion plates short, teeth irregular, slit rays clearly visible in head and intermediate valves, scarcely visible in tail valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0C4EF40FADF9DF6B59921D.taxon	discussion	Remarks. This species was initially attributed to the genus Tonicella Carpenter, 1873 by Dell’Angelo et al. (2020 b), and later included by Sirenko & Dell’Angelo (2023) in the genus Lepidochitona Gray, 1821. There is a certain variability in the shape of intermediate valves (the width / length ratio ranges from 2.05 to 2.50, and the anterior profile, with H / W = 0.41 – 0.51) and tail valves (the width / length ratio ranges from 1.93 to 2.36). The head valves have a sculpture with slight presence of radial folds. Comparisons. Lepidochitona nuda (Dell’Angelo, Lesport, Cluzaud & Sosso, 2020) is similar to L. tenuissima (Sandberger, 1859) (see below), from which it differs mainly by the smooth surface with the presence of poorly or hardly visible growth lines (very prominent along the entire surface of the valves in L. tenuissima), and by the shape of intermediate valves (W / L = 2.05 – 2.50 vs. 2.41 – 2.96 in L. tenuissima; H / W = 0.41 – 0.51 vs. 0.39 – 0.42 in L. tenuissima).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0C4EF40FADF9DF6B59921D.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0B4EF60FADF8E06860919C.taxon	description	Fig. 117	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0B4EF60FADF8E06860919C.taxon	materials_examined	Type material. Syntypes: NHMW 1858 / 0018 / 0255, 47 valves, Figs 117 A – D; NHMW 2011 / 0009 / 0006, 2 valves. Type locality. Gaas, Larrat (France). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene (Rupelian): Gaas: type material (syntypes) plus Benoist collection MHNBx [MHNBx 2014.14.32.1 – 2014.14.32.3)]; ZSM Mol 20060781 [12 valves (described as Stenoplax monila Cherns & Schwabe, 2019)]; Rupelian levels: 208 valves (BD 941, Figs 117 E – L, MZB 50569 – 50570); Espibos: 45 valves (AC, DA, PR); Lagouarde: 56 valves (AC, DA); Larrat: 3,289 valves (AC, Figs 117 O – P, DA, Figs 117 M – N, JCV, MHNBx 2020.2.1 – 2020.2.3). Maximum width of the valves: 3.8 / 5.5 / 4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0B4EF60FADF8E06860919C.taxon	description	Description. Head valve semicircular, some radial folds (6 – 8) may be present. Intermediate valve broadly rectangular (W / L = 2.38 – 3.44), rounded in anterior profile, moderately to highly elevated (H / W = 0.27 – 0.54), anterior margin almost straight or slightly sinuose, slightly concave in the jugum, side margins weakly rounded, posterior margin straight, apex not evident or just highlighted, lateral areas scarcely differentiated. Tail valve semicircular to triangular, W / L = 1.41 – 1.77, anterior margin slightly sinuose, slightly concave in jugum, mucro subcentral, antemucronal and postmucronal slopes almost straight, slightly concave behind mucro. Tegmentum entirely covered by close-set, irregularly arranged granules, some coalescent, roundish in head valves (diameter ca 40 – 45 μm), more irregular and tending to rhomboidal in other valves (maximum length ca 50 – 68 μm). Each granule with 8 – 12 aesthetes of equal size, often one (or more) of them centrally located. Articulamentum with rounded apophyses, separated by a wide jugal sinus, slit formula: 10 – 11 / 2 / 9 – 13, slits inequidistant, the second slit in intermediate valves lies close to posterior margin, slit rays clearly visible in head and intermediate valves, teeth short.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0B4EF60FADF8E06860919C.taxon	discussion	Remarks. A study of a large quantity of valves from Gaas was carried out by Dell’Angelo et al. (2020 b), who also examined the valves preserved in the Benoist collection (MHNBx 2014.14.32.1 – 2014.14.32.3), attributed to Lepidochitona oligocaena (Rolle, 1862). In addition, these Authors examined the valves preserved at ZSM as Stenoplax monila Cherns & Schwabe (2019), concluding that this species is a younger synonym of L. oligocaena. Lepidochitona oligocaena is characterized by a remarkable morphological variability (Dell’Angelo et al., 2020 b). Some intermediate valves (Figs 117 M – N) are more elongated (W / L = 2.91 – 3.44) and lower in anterior profile (H / W = 0.27 – 0.32), but the sculpture is similar with the same rhomboidal granules of L. oligocaena with aesthetes of equal size, and these valves have been considered to fall within the range of variability of L. oligocaena. The same for some tail valves (Figs 117 O – P) more than semicircular or tending to a triangular shape, compare Figs 117 J and 113 O. Comparisons. Lepidochitona oligocaena is characterized by the intermediate valves showing a posterior margin straight with the apex not evident or just highlighted, LA scarcely differentiated and the tegmentum covered by irregular subrhomboidal granules with aesthetes of equal size, not differentiated. The only other species of Lepidochitona from Gaas with similar characters is L. tessellata Dell’Angelo, Lesport, Cluzaud & Sosso, 2020, which however differs in having different and coarser sculpture on LA and PMA (see below).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF0B4EF60FADF8E06860919C.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Rolle, 1862; Dell’Angelo et al. 2019 b, 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF094EF70FADFB60685E900D.taxon	description	Fig. 118	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF094EF70FADFB60685E900D.taxon	materials_examined	Type material. Holotype: MNHN. F. A 67131, intermediate valve, width 3.6 mm, Figs 118 A – C. Paratypes: MNHN. F. A 67134 – F. A 67136 (3 valves), Figs 118 D, 118 G – H; NHMW 2017 / 0108 / 0038 – 2017 / 0108 / 0040 (3 valves); RGM. 1008360 – 1008363 (4 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): Saint-Clément-de-la-Place: type material plus 136 valves (MNHN. F. A 67137, Figs 118 E – F, MNHN. F. A 67138, NHMW 2017 / 0108 / 0041, RGM. 1008356, RGM. 1008364, RGM. 1008436, BD 152). Maximum width of the valves: 2.8 / 3.6 / 2.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF094EF70FADFB60685E900D.taxon	description	Description. Head valve semicircular, with few radial ridges that correspond to slits on insertion lamina. Intermediate valve broadly rectangular (W / L = 2.40 – 2.65), semicarinate in anterior profile, moderately elevated (H / W = 0.40 – 0.47, 0.40 in holotype), side margins rounded, posterior margin almost straight on both sides of prominent apex, lateral areas strongly delimited from central area by diagonal fold, second weaker diagonal fold present on posterior margin. Tail valve elliptical (W / L = 2.08 – 2.23), anterior margin straight, mucro subcentral, antemucronal slope straight to slightly convex, postmucronal slope concave. Tegmentum smooth and glossy, with several prominent growth lines, aesthetes of same size. Articulamentum with apophyses not well preserved, wide, short, anteriorly rounded, separated by narrow straight sinus, insertion plates short, slit formula 9 / 1 / 7 – 8 (a second slit ray lies close to posterior margin on intermediate valves), teeth irregular, slightly thickened at edges, slit rays conspicuous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF094EF70FADFB60685E900D.taxon	discussion	Remarks. This species was initially attributed to the genus Tonicella Carpenter, 1873 by Dell’Angelo et al. (2018 b), and later included by Sirenko & Dell’Angelo (2023) in the genus Lepidochitona Gray, 1821. The valves are small and not well preserved, only a few are complete, showing little intraspecific variability; some intermediate valves are more elevated (H / W = 0.47) and more rounded in profile (Figs 118 E – F). Comparisons. Tonicella marmorea (Fabricius, 1780) and Boreochiton ruber (Linnaeus, 1767) differ from Lepidochitona redoniensis (Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2028) by lacking radial ridges on the head valve and the diagonal fold separating lateral and central areas on the intermediate valves and being larger.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF094EF70FADFB60685E900D.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Ligerian Basin, France: Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF084EF90FADFA13686094C8.taxon	description	Fig. 119	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF084EF90FADFA13686094C8.taxon	description	1934, p. 4]. non Chiton reussi Rzehak, 1893; Rzehak, 1893, p. 171 (an undeterminable species, considered by Rzehak = Chiton siculus? Reuss).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF084EF90FADFA13686094C8.taxon	materials_examined	Type material. Syntypes: NHMW 2011 / 0009 / 0002, 1 tail valve; NHMW 2011 / 0009 / 0003, 4 valves (3 intermediate and 1 tail), Figs 119 A – E; NHMW 2011 / 0009 / 0004, 1 intermediate valve; NHMW 2011 / 0009 / 0008: 5 intermediate valves (Dell’Angelo et al., 2019 b, figs 5 A – F). Type locality. Gaas (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas: Syntypes NHMW plus Rupelian levels: 9 valves (BD 942, Fig. 119 I); Espibos: 3 valves (AC, DA, PR); Lagouarde: 3 valves (DA); Larrat: 209 valves (AC, DA, JCV, Figs 119 J – L, MHNBx 2020.1.1, MHNBx 2020.3.4, Figs 119 F – H, MHNBx 2020.3.5, MHNBx 2020.1.7). Maximum width of the valves: 4.1 / 5.3 / 2.7 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF084EF90FADFA13686094C8.taxon	description	Description. Head valve semicircular, weak elevated, some radial fold may be present, slope straight. Intermediate valve broadly rectangular (W / L = 2.45 – 2.83), carinate in anterior profile, moderately elevated (H / W = 0.28 – 0.33), anterior margin straight or slightly concave, lateral margins rounded, posterior margin slightly concave on both sides of prominent apex, lateral areas separated from central areas by a diagonal fold. Tail valve semicircular (W / L = 1.60 – 2.06), anterior margin slightly concave, mucro subcentral, acute, antemucronal slope straight, postmucronal slope concave. Tegmentum entirely covered by irregularly arranged and very close-set roundish granules, not elevated, some coalescent, diameter up to 78 μm in HV, 83 μm in intermediate valves, 96 μm in tail ones; each granule bearing a central megalaesthete, and 6 – 10 micraesthetes arranged irregularly along margin, additional micraesthetes randomly placed between granules. Articulamentum with small triangular apophyses in intermediate valves, trapezoidal in tail valve, slit formula 8 – 9 / 2 / 8 – 9, slits deep, teeth strong, very uneven in width, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF084EF90FADFA13686094C8.taxon	discussion	Remarks. De Rochebrune (1882: p. 57) considered “ Chiton reussi ” a synonym of “ Chiton oligocaenus Rolle, 1862 ”; however, the study of the type material of Lepidochitona reussi (Rolle, 1862) at NHMW clarified that the two species are, in fact, distinct (Dell’Angelo et al. 2019 b.) The studied material is generally well preserved, many valves (but not all) show a truncation in the posterior margin (Figs 119 J – L). The valves figured by Varone (2008: figs 13 – 14) as Chiton sp. 1 show this characteristic truncation. Comparisons. Lepidochitona reussi (Rolle, 1862) differs from L. oligocaena (Rolle, 1862) by the shape of intermediate valves (carinate with H / W = 0.28 – 0.33 vs. rounded with H / W = 0.27 – 0.54 in L. oligocaena), the larger granules (roundish with diameter up to 96 μm vs. subrhomboidal with L up to 68 μm in L. oligocaena), the structure of the aesthetes (with one central megalaesthete and 6 – 10 micraesthetes vs. aesthetes of equal size in L. oligocaena), and the number of slits in the tail valves (8 – 9 vs. 9 – 13 in L. oligocaena).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF084EF90FADFA13686094C8.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Rolle 1862; Dell’Angelo et al. 2019 b, 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF064EFA0FADFDAB6A949490.taxon	description	Fig. 120	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF064EFA0FADFDAB6A949490.taxon	materials_examined	Type material. Holotype RGM. 1364017, intermediate valve, width 2.3 mm (Figs 120 A – D). Paratype RGM. 1364018, intermediate valve (width 4.8 mm). Type locality. Vale de Freixo site, near the village of Carnide, Pombal municipality (central-west Portugal). Type stage. Pliocene, upper Zanclean to lower Piacenzian, Carnide Formation, basal fossiliferous grey sands, “ Bed 3 ” in Gili et al. (1995) and Dell’Angelo & Silva (2003). Material examined. Pliocene: Portugal: Vale de Freixo: type material plus 14 intermediate valves: (BD 245, GeoFCUL VFX. 03.361, Figs 120 E – H, GeoFCUL VFX. 03.344 – VFX. 03.345, RGM. 1364019, MNHN. F. A 81991). Maximum width 4.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF064EFA0FADFDAB6A949490.taxon	description	Description. Head and tail valves unknown. Intermediate valves broadly rectangular (W / L = 2.70 – 2.90), elevated (H / W = 0.40 – 0.50), carinate in anterior profile, anterior margin convex with jugal part almost straight, side margins slightly rounded, posterior margin straight or slightly concave on both sides of large and prominent apex, lateral areas slightly raised, sharply delimited. LA sculptured with fine elongate granules, coalescing into continuous, radially oriented lines, few concentric growth lines more evident towards side margins. CA covered for about 60 % of valve width by roundish irregular granules, almost all separated, but some coalescent, with up to eight aesthetes of equal size; 40 % continuing towards side margins by obliquely oriented and few concentric growth lines joined with those present on lateral areas. Each granule with up to 8 aesthetes of equal size in CA. Articulamentum with poorly preserved apophyses, round, apical area triangular, insertion plate short with two slits, slit ray well visible (second slit ray lies close to posterior margin).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF064EFA0FADFDAB6A949490.taxon	discussion	Remarks. The fossil record of Lepidochitona rochae Dell’Angelo, Landau, Silva & Sosso, 2022 is limited to the Pliocene of Portugal, based upon of a few intermediate valves, incomplete and poorly preserved. However the generic attribution has ben problematic, and the species was assigned to Lepidochitona mainly based on the similarity of the granules in the central area of the valves with the sculpture typical of this genus. Comparisons. Lepidochitona rochae is well characterized by the granules of the tegmental sculpture covering only a part of the whole valve.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF064EFA0FADFDAB6A949490.taxon	distribution	Distribution. Pliocene: Northeastern Atlantic, Mondego Basin, Portugal: Vale de Freixo (Dell’Angelo et al. 2022).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF054EFB0FADFE6C6B599604.taxon	description	Fig. 121	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF054EFB0FADFE6C6B599604.taxon	materials_examined	Type material. Holotype MHNBx 2020.2.4, intermediate valve, width 3.2 mm, Figs 121 G – I. Paratypes: MHNBx 2020.1.2, head valve; MHNBx 2020.2.5, tail valve, Figs 121 J – L. Type locality. Gaas, Larrat (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas: type material plus MHNBx Benoist collection (MHNBx 2014.14.23.1 – 2014.14.23.4, labeled Tonicia gaasensis, Figs 121 A – C); Rupelian levels: 17 valves (BD 943); Espibos: 8 valves (AC, DA); Lagouarde: 2 valves (AC); Larrat: 323 valves (AC, DA, JCV, MHNBx 2020.1.2, MHNBx 2020.2.4 – 2020.2.5, MHNBx 2020.1.8, Figs 121 D – F). Maximum width of the valves: 2.6 / 4.2 / 2.6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF054EFB0FADFE6C6B599604.taxon	description	Description. Head valve semicircular, elevated, some radial fold may be present, slope slightly convex. Intermediate valve broadly rectangular (W / L = 1.83 – 2.26), semicarinate in anterior profile, moderately elevated (H / W = 0.39 – 0.47), anterior margin straight, lateral margins slightly rounded, posterior margin slightly concave at both sides of prominent apex, lateral areas separated from central area by a diagonal fold. Tail valve triangular (W / L = 1.33 – 1.38), anterior margin straight, mucro subcentral, antemucronal and postmucronal slopes straight, almost aligned. Tegmentum entirely covered by irregularly arranged, elevated, roundish granules, some coalescent, diameter up to 100 μm in head valve, tending to be more oval near anterior margin, 126 μm in intermediate valves, 100 μm in tail ones; each granule with a subcentral megalaesthete, and up to 10 micraesthetes arranged irregularly along margin, additional micraesthetes randomly placed between granules. Articulamentum with large apophyses, slit formula 9 / 2 / 9, slit rays clearly visible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF054EFB0FADFE6C6B599604.taxon	discussion	Remarks. The four valves from the Benoist collection at MHNBx (from MHNBx 2014.14.23.1 to MHNBx 2014.14.23.4), labeled Tonicia gaasensis de Rochebrune, 1882, have been considered conspecific to Lepidochitona tarbelliana Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 by Dell’Angelo et al. (2020 b). Comparisons. This species has larger granules than those of Lepidochitona oligocaena (Rolle, 1862) and L. reussi (Rolle, 1862), the tail valves, similar to those of L. reussi, differ, in fact, for their shape (W / L = 1.33 – 1.38 vs. 1.60 – 2.06 of L. reussi) and in the frontal view of the valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF054EFB0FADFE6C6B599604.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF044EFB0FADFC186B5992C8.taxon	description	Fig. 122	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF044EFB0FADFC186B5992C8.taxon	materials_examined	Type material. Holotype MHNBx 2020.3.1, tail valve, width 4.3 mm, Figs 122 I – L. Paratypes: MHNBx 2020.3.2, head valve, Figs 122 A – C; MHNBx 2020.3.3, intermediate valve, Figs 122 E – H. Type locality. Gaas, Larrat (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas: type material plus Rupelian levels: 6 valves (BD 944); Espibos: 25 valves (AC, DA, PR); Lagouarde: 6 valves (AC, DA, PR); Larrat: 119 valves (AC, DA, JCV, Fig. 122 D). Maximum width of the valves: 4.1 / 7 / 5.1 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF044EFB0FADFC186B5992C8.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, slope almost straight. Intermediate valve broadly rectangular (W / L = 2.17 – 2.48), rounded in anterior profile, moderately elevated (H / W = 0.41), anterior margin almost straight, lateral margins rounded, posterior margin almost straight, apex not evident, lateral areas evidenced by different sculpture. Tail valve semicircular (W / L = 1.38 – 1.52), anterior margin straight, mucro subcentral, antemucronal slope slightly convex, postmucronal slope slightly concave just behind mucro. Tegmentum entirely covered by randomly arranged granules, subrhomboidal in CA and AMA (length 90 – 100 μm); smaller, roundish in LA and PMA (length ca. 50 μm), some higher tending to form elevated growth lines or groups of isolated granules (some coalescent), giving the impression of forming rough surface. Each granule with many aesthetes of equal size, up to 15 in more elongate granules in CA and AMA, up to 10 in roundish granules in LA and PMA, often one (or more) of them centrally located. Articulamentum with large apophyses, teeth inequidistant, slit formula 9 – 11 / 2 / 9 – 11, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF044EFB0FADFC186B5992C8.taxon	discussion	Remarks. Our material is well preserved, revealing the coarse sculpture of the tegmentum, characteristic for this species. Comparisons. This species is superficially similar to Lepidochitona oligocaena (Rolle, 1862), from which it differs mainly in the different sculpture of LA and PMA, very irregular in L. tessellata Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 with the presence of elevated granules forming growth lines or irregularly distributed groups of higher granules, while in L. oligocaena the granules in these areas are regularly distributed, without any trace of growth lines.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF044EFB0FADFC186B5992C8.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF034EFD0FADFB546B599104.taxon	description	Fig. 123	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF034EFD0FADFB546B599104.taxon	materials_examined	Type material. Holotype MHNBx 2020.3.7, an intermediate valve, width 2.9 mm, Figs 123 A – D. Type locality. Gaas, Larrat (France). Type stage. Oligocene (Rupelian). Material examined. Oligocene (Rupelian): Gaas (Larrat): type material plus 1 intermediate valve, width 3.5 mm (JCV).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF034EFD0FADFB546B599104.taxon	description	Description. Head and tail valves not present. Intermediate valve rectangular (W / L = 1.68), rounded in anterior profile, highly elevated (H / W = 0.47 – 0.56), anterior margin weakly sinuose, lateral margins slightly rounded, posterior margin almost straight at both sides of poorly developed apex, lateral areas not raised, but separated from central area by a diagonal fold. Tegmentum entirely covered by irregularly arranged, weak elevated, roundish granules, not coalescent, diameter up to 100 μm; each granule bearing a subcentral megalaesthete, and up to 6 micraesthetes arranged irregularly along margin, additional micraesthetes randomly placed between granules. Articulamentum with rounded apophyses, slit formula two slits for each side, slit rays visible, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF034EFD0FADFB546B599104.taxon	discussion	Remarks. This species is characterized by the tegmentum entirely covered by roundish granules (Fig. 123 D), the “ enveloping ” valve with a characteristic rounded and high anterior profile (Fig. 123 C), and an almost straight posterior margin with a small apex (Fig. 123 A). A second lower intermediate valve (H / W = 0.47, see Dell’Angelo et al. 2020 b: fig. 15. E – F) has a different shape (W / L = 1.95), but the sculpture and the posterior straight margin with the small apex are similar, and has been considered falling within the range of variability of L. vancuycki Dell’Angelo, Lesport, Cluzaud & Sosso, 2020. Comparisons. The closest species is Lepidochitona charlesi Dell’Angelo, Lesport, Cluzaud & Sosso, 2020, from which L. vancuycki Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 differs by the almost straight posterior margin with a small apex (vs. slightly concave at both sides of the prominent apex in L. charlesi), LA not raised, but separated from CA by a diagonal fold (vs. scarcely differentiated in L. charlesi), and the smaller roundish granules (up to 100 μm, vs. triangular-oval up to 110 μm diameter in L. charlesi).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF034EFD0FADFB546B599104.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF024EFD0FADFB186FE5928B.taxon	description	and unpublished observations). ...... continued on the next page	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF014EE00FADF90B6B69947C.taxon	description	Fig. 124	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF014EE00FADF90B6B69947C.taxon	materials_examined	Type material. Holotype: MZ VIII Ma- 65, head valve (Fig. 124 A). Type locality. Węglinek (Poland). Type stage. Middle Miocene. Material examined. Middle Miocene: Eastern Paratethys: Ukraine: Varovtsi: 3 valves (BD 945, Figs 124 J – L), Velyka Levada: 2 valves (BD 946, Figs 124 D – I). Maximum width of the valves: 1.3 / 2.6 / 2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF014EE00FADF90B6B69947C.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular (W / L = 2.40 – 2.75), moderately elevated (H / W = 0.33 – 0.40), carinate in anterior profile, anterior margin almost straight to slightly convex, side margins rounded, posterior margin almost straight, with well pronounced apex, lateral areas slightly raised. Tail valve semicircular (W / L = 1.78), mucro subcentral, antemucronal slope slightly convex, postmucronal slope slightly convex. Tegmentum uniformly covered with coarse granules of irregular shape, randomly arranged and distinctly separated, which appear radially oriented in HV and PMA, 60 – 80 µm long in CA, each granule with a central megalaesthete, and up to 6 – 7 micraesthetes irregularly distributed. Articulamentum with apophyses wide, triangular, slit formula 8 / 1 / 8 – 11, teeth slightly roughened on outside, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF014EE00FADF90B6B69947C.taxon	discussion	Remarks. Lepidochitona baluki Macioszcsyk, 1988 was described upon four valves (1 head, 2 intermediate and 1 tail) of small size (maximum width 1.5 mm) from Węglinek (Poland) (Figs 124 A – C) and has never been found after. We attribute to Lepidochitona baluki two valves from Velyka Levada and 3 from Varovtsi, which have a tegmentum covered with coarse oval granules, and a similar shape. Macioszcsyk (1988) described also a single tail valve, considerably arched from Węglin as Lepidochitona sp. This valve is closest to L. baluki, in the tegmentum’s sculpture, it is greater (width 2.7 mm vs. 1.4 in L. baluki), and differs mainly for the number of slits, 11 vs. 8 in L. baluki. We consider for the moment Lepidochitona sp. falling within the variability of L. baluki, pending the availability of further material that could confirm or not its specific attribution. The two intermediate valves from Bánd (Hungary), described by Dulai (2005) as Lepidochitona sp., could be attributable to L. baluki. Comparisons. The sculpture of the tegmentum is coarser than the three other species of Lepidochitona known from the Paratethys. See Tab. 20 for more comparison with the other Lepidochitona spp. from the Paratethys considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF014EE00FADF90B6B69947C.taxon	distribution	Distribution. Middle Miocene. Central Paratethys (Langhian-Serravallian): Poland: Węglin, Węglinek (Macioszczyk 1988), Hungary: Bánd (Dulai 2005; this study); Eastern Paratehys: Ukraine: Varovtsi, Velyka Levada (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1F4EE20FADFE006F5C9044.taxon	description	Fig. 125	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1F4EE20FADFE006F5C9044.taxon	materials_examined	Type material. Unknown, not present in NHMV. Type locality. Rudoltice (Czech Republic). Type stage. Middle Miocene. Material examined. Middle Miocene: Central Paratethys: Romania: Lăpugiu de Sus: 1 valve (NHMV 2010 / 0256 / 0017), Kostej: 2 valves (NHMV 2010 / 0256 / 0018, Figs 125 E – H, NHMV 2010 / 0256 / 0021, Figs 125 M – P); Eastern Paratehys: Ukraine: Varovtsi: 120 + valves (BD 947, Figs 125 A – C, 125 I – L); Hungary: Letkés: 2 valves (BD 948), Várpalota: 2 valves (BD 949, Fig. 125 D). Maximum width of the valves: 3.5 – 5.2 – 2.1 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1F4EE20FADFE006F5C9044.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, with or without presence of radial depressions. Intermediate valves broadly rectangular (W / L = 2.40 – 2.70), moderately elevated (H / W = 0.30 – 0.43), semicarinate in anterior profile, anterior margin almost straight to slightly concave, side margins rounded, posterior margin almost straight, with strongly protruded apex, lateral areas slightly raised, not distinctly separated from central area. Tail valve semicircular (W / L = 1.88 – 2.12), anterior margin almost straight, mucro in variable position, from anterior to posterior, anterior margin almost straight, antemucronal slope straight, postmucronal slope slightly concave. Tegmentum uniformly covered with fine roundish / oval flat granules irregularly arranged, which give the impression of being arranged in two systems of rows, running irregularly from apex and posterior margin towards anterior margin, almost forming a net-like ornament. Each granule up to 80 µm long, with a central megalaesthete and 7 – 10 micraesthetes along margin. Articulamentum with apophyses wide, triangular in intermediate valves, quadrangular in tail valve, slit formula 8 – 10 / 1 / 12, teeth massive, slightly roughened on outside, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1F4EE20FADFE006F5C9044.taxon	discussion	Remarks. The taxon Chiton lepidus Reuss, 1860 is preoccupied by Chiton lepidus Gould, 1859, now considered as a synonym of Lepidozona luzonica (Sowerby, 1842), an Indo-Pacific species from the Philippines to the Arabian Gulf. Schwabe (2000), the first author noticing this homonymy, proposed to maintain the specific name of Chiton lepidus Reuss, 1860 for a chiton (currently known as Lepidochitona lepida) from the Middle Miocene of Europe. The International Commission on Zoological Nomenclature (2003) has accepted this proposal (Opinion 2033). Chiton lepidus was erected by Reuss (1860) based upon an intermediate valve from the Middle Miocene of Rudoltice in the Moravian part of the Carpathian Foredeep Basin, Czech territory. Reuss’ original material is no longer available. Šulc (1934) described and later illustrated a head and a tail valve from the same locality, attributed to this species. The paucity of material and the opinability of intepreting species in Lepidochitona may accounts for the conservative approach by later authors who synonymized C. lepidus with Mediterranean extant taxa; L. caprearum (Scacchi, 1836) by Malatesta (1962) and Laghi (1977); L. cinerea (Linnaeus, 1767) by Šulc [1934, = Lepidopleurus marginatus (Pennant) reported by Sacco 1897], Bałuk (1971), and Laghi (1977); L. monterosatoi Kaas & Van Belle, 1981 by Bałuk (1984); L. canariensis (Thiele, 1909) by Studencka & Studencki (1988). For example, Macioszczyk (1988) considered the valves of L. lepida figured by Bałuk (1971: pl. 4, figs 6 – 12) as belonging to L. monterosatoi (pl. 4, figs 6, 8, 10) and L. caprearum (pl. 4, fig. 9), while Studencka & Studencki (1988: p. 40) considered the tail valve figured by Bałuk (1971: pl. 4, figs 11 – 12) not corresponding to the tail valve of L. lepida figured by Šulc (1934: pl. 1, fig. 15). The inadequacy of the descriptions of Reuss and Šulc, and the scarcity of records in literature (sometimes attributed to different species) prompted our reconsideration of the available material, focusing upon some morphological characters: - The presence of radial depressions on the head valve is not evidenced in the Šulc’s description, although very slight radial depressions are observable on the head valve (Šulc 1934: pl. 1, fig. 13). Literature records figure valves with (Bałuk 1971: pl. 4, fig. 6) and without (Studencka & Studencki 1988: pl. 2, fig. 1) radial depressions, a situation observed also in our own material both (Figs 125 A, 125 D). - The evaluation of the tail valve is open to interpretation: Šulc’s description reports the mucro in posterior position, at two-thirds the length, while the few tail valves in our material from Varovtsi (Ukraine) and Kostej (Romania) show a mucro slightly in anterior or at most subcentral position (Figs 125 L, 125 P). In literature, only Bałuk (1971: pl. 4, figs 11 – 12) show tail valves from Korytnica (Poland) with the mucro in posterior position, while the only other tail valve figured (Bałuk 1984: pl. 7, fig. 3) show the mucro in anterior or at most subcentral position. For these reasons we consider attributable to Lepidochitona lepida tail valves with the mucro in a wide range of positions. Therefore, we hypothesize that also the 19 valves (3 head and 16 intermediate) from Węglinek cited by Macioszcsyk (1988) as Lepidochitona monterosatoi Kaas & Van Belle, 1981, may well fall within the variability of L. lepida. Šulc (1934) described another species of Lepidochitona (as Middendorffia boravičensis) upon a single intermediate valve (width 3.3 mm) from Borač (Czech Republic); this taxon differs from L. lepida basically by its slightly different shape (3.3 x 1.2 mm vs. 4 x 1.1 mm in L. lepida) and lack of a longitudinal keel, we judge that such features are of little taxonomic significance, and thus consider Lepidochitona boravičensis as a junior synonym of L. lepida. Comparisons. The closest species is Lepidochitona kieli nov. sp (see below).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1F4EE20FADFE006F5C9044.taxon	distribution	Distribution. Middle Miocene. Central Paratethys (Langhian – Serravallian): Slovakia: Devínska Nová Ves, Dubová, Rohožník, (Ruman & Hudáčková 2015), Czech Republich: Borač, Rudoltice (Reuss 1860; Šulc 1934), Poland: Korytnica, Lychów, Nawodzice, Niskowa, Rybnica, Węglin, Węglinek (Bałuk 1965, 1971, 1984; Macioszczyk 1988; Studencka & Studencki 1988), Romania: Kostej, Lăpugiu de Sus (Šulc 1934; this study), Hungary: borehole Szokolya- 2, Devecser, Hidas, Letkés, Várpalota (Csepreghy-Meznerics 1950; Dulai 2001, 2025 a; Dulai & Katona 2024; this study); Eastern Paratehys: Ukraine: Podhorce, Varovtsi (Studencka & Dulai 2010; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	description	Fig. 126	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	materials_examined	Type material. Holotype MSNG 62665, intermediate valve, width 5.5 mm (Figs 126 A – E). Paratype 1: MSNG 62666, tail valve, width 2.1 (Figs 126 K – L). Paratype 2: MNHN. F. A 98478, intermediate valve, width 3 mm (Figs 126 F – H). Type locality. Varovtsi (Ukraine). Type stage. Middle Miocene.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	etymology	Etymology: The species is named after Steffen Kiel in recognition of his outstanding contribution to the taxonomy and biogeography of fossil molluscs.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	materials_examined	Material examined. Middle Miocene: Eastern Paratethys: Ukraine: Varovtsi: type material plus 8 valves (BD 950, Figs 126 I – J). Maximum width of the valves: 1.8 / 5.5 / 2.1 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	diagnosis	Diagnosis. Head valve semicircular, intermediate valves broadly rectangular, moderately elevated, semicarinate, apex strongly protruded, tail valve semicircular, mucro in anterior position. Tegmentum uniformly covered with flat granules, irregularly arranged, elliptical in CA, AMA, more roundish and smaller in HV, LA, PMA. Articulamentum with apophyses wide, triangular, teeth massive, one slit in intermediate valves	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, without presence of radial depressions. Intermediate valves broadly rectangular (W / L = 2.40 – 2.79), moderately elevated (H / W = 0.36 – 0.47), semicarinate in anterior profile, anterior margin almost straight, side margins rounded, posterior margin almost straight, with strongly protruded apex, lateral areas slightly raised, not distinctly separated from central area. Tail valve semicircular (W / L = 1.95), anterior margin slightly concave, mucro in anterior position. Tegmentum uniformly covered with flat granules, irregularly arranged, elliptical up to 125 µm long in CA and AMA, more roundish and smaller in HV, LA, PMA. Each granule with a subcentral megalaesthete and up to 10 micraesthetes irregularly disposed. Articulamentum with apophyses wide, triangular, one slit in intermediate valves, teeth massive, slightly roughened on outside, eaves spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	discussion	Remarks. Some intermediate valves from Varovtsi (Ukraine) display morphological characteristics to justify the establishment of a new species. Comparisons. The closest species is Lepidochitona lepida (Reuss, 1860), from which L. kieli sp. nov. differs mainly for the different sculpture, with larger elliptical granules (up to 125 µm long) most closely spaced, vs granules more roundish and smaller (up to 80 µm long), and a different structure of aesthetes.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1D4EE30FADFA586BF69204.taxon	distribution	Distribution. Middle Miocene. Central Paratethys (Langhian-Serravallian): Eastern Paratehys: Ukraine: Varovtsi (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1B4EE40FADFF05694692C6.taxon	description	Fig. 127	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1B4EE40FADFF05694692C6.taxon	materials_examined	Type material. Holotype BkK-A 26 (Bałuk collection), tail valve (Fig. 127 A). Type locality. Korytnica, Holy Cross Mts. (Poland). Type stage. Middle Miocene. Material examined. No actual material available, only descriptions and illustrations from the literature.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1B4EE40FADFF05694692C6.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped. Intermediate valves broadly rectangular, anterior margin almost straight, side margins rounded, posterior margin almost straight, with well pronounced apex, lateral areas slightly raised. Tail valve triangular, flat, anterior margin almost straight, mucro subcentral. Tegmentum uniformly covered with closely spaced, fine granules, arranged in intersecting, arcuate rows. Articulamentum with teeth roughened on outside, slit formula 8 / 1 / 7, slit rays visible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1B4EE40FADFF05694692C6.taxon	discussion	Remarks. Lepidochitona subgranosa Bałuk, 1971 is known only from the Middle Miocene of Poland, Korytnica (Bałuk 1971, 1984), Nawodzice (Figs 127 C – D) and Rybnica (Studencka & Studencki 1988). The sculpture of the tegmentum is similar to that of Lepidochitona lepida (Reuss, 1860), the only difference is represented by the tail valve (triangular, flat, mucro subcentral, vs. semicircular, more elevated, mucro anterior to posterior in L. lepida) [Bałuk 1971, p. 460: “ Much less prominent granules and, primarily, a different shape and degree of convexity of the tail valve differ them from Lepidochitona lepida ”; Studencka & Studencki 1988, p. 40: “ The ornamentation of tegmentum in L. subgranosa resembles the sculpture of L. lepida, but the latter species has longer, triangular tail valve with posterior mucro, and 8 radial ribs at its head valve (in adult specimens) ”]. The tail valve figured by Studencka & Studencki 1988 (Fig. 127 D) shows the same triangular shape of the holotype of L. subgranosa, and so we confirm for the moment the attribution to the the latter species, pending the discovery of further material, also because there are no indications that the valve is “ flat ” as indicated (but not figured) in the original description Laghi (1977) postulated that both Lepidochitona lepida and L. subgranosa are conspecific with the Recent species L. cinerea (Linnaeus, 1767), a hypothesis rejected by Bałuk (1984) as this cannot be evidenced when having only isolated valves. On the other hand Bałuk himself (1984) highlighted the remarkable similarity of the sculpture of the valves of this species with Lepidochitona canariensis (Thiele, 1909), and in the material from Korytnica there are also some plates “ with an ornamentation particularly fine and these specimens are fairly distinctly different than the rest ” (Bałuk 1971: 460). Comparisons. See Tab. 20 for further comparison with the other Lepidochitona spp. from the Paratethys considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF1B4EE40FADFF05694692C6.taxon	distribution	Distribution. Middle Miocene. Central Paratethys (Langhian – Serravallian): Poland: Korytnica, Nawodzice, Rybnica (Bałuk 1971, 1984; Studencka & Studencki 1988).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF194EE70FADFDA569C593A0.taxon	description	Fig. 128	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF194EE70FADFDA569C593A0.taxon	materials_examined	Type material. Lectotype SMF 14.32 a designated by Janssen 1978, the intermediate valve figured by Boettger (1969: pl. 1, figs 14 a – 14 c); Paralectotype SMF 14.32 b. Type locality. Gienberg (Germany). Type stage. Oligocene (Rupelian). Material examined. Oligocene: France: Ormoy-La-Rivière: 60 valves (BD 951, Figs 128 A – L); Auvers-Saint-Georges: (3 valves (BD 952); Morigny: 2 valves (BD 953). Maximum width of the valves: 3.4 – 4.3 – 1.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF194EE70FADFDA569C593A0.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, some radial folds may be present. Intermediate valves broadly rectangular (W / L = 2.14 – 2.75), moderately to highly elevated (H / W = 0.38 – 0.60), semicarinate in anterior profile, anterior margin almost straight to slightly concave between apophyses, side margins rounded, posterior margin almost straight, with a small apex, lateral areas from scarcely differentiated from central areas to raised and separated by a diagonal fold. Tail valve semicircular, elevated, anterior margin almost straight, mucro subcentral or slightly anterior, AMA and PMA areas poorly differentiated, postmucronal slope slightly concave. Tegmentum uniformly covered with rounded to diamond-shaped granules, irregularly arranged in more or less distinct vertical rows in CA and AMA, in diverging rows running obliquely in LA. Articulamentum with apophyses wide, but only slightly projecting, with broad sinus, slit formula 8 / 2 / 8, the second small slit in intermediate close to posterior margin, eaves coarsely spongy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF194EE70FADFDA569C593A0.taxon	discussion	Remarks. This species was described by Boettger (1869) upon two intermediate valves from the Oligocene of Gienberg (Germany). Later, Janssen (1978) re-examined the type material and considered Tonicia etrechyensis de Rochebrune, 1882 (nom. nov. pro Chiton terquemi Deshayes, 1861, non C. terquemii Deslongchamps, 1859, from the lower Jurassic of France) and Tonicia mamillata de Rochebrune, 1882, identical to Lepidochitona corrugis (Boettger, 1869), therefore junior synonyms of the latter species. Janssen (1978) figured two intermediate valves from Ormoy-La-Rivière (type locality of T. mamillata) and two valves (intermediate and tail) from Auvers-Saint-Georges. All three species (Lepidochitona corrugis, L. etrechyensis, L. mamillata) have been described from intermediate valves, with the first figure of a tail valve provided by Janssen (1978: pl. 16 fig. 43). Boettger evidenced the strong similarity of C. corrugis with C. terquemi Deshayes, 1861. The taxa could be distinguished by the greater elevation of the valves: 0.35 (C. corrugis) vs. 0.48 (C. terquemi), extracted from Boettger and Deshayes’ figures, respectively for), and the subdued lateral areas, without a diagonal rib to separate LA from CA in C. terquemi. The abundant material (Ormoy-La-Rivière and Auvers-Saint-Georges) available to study documents the great morpological variability of some characters, e. g.,: (1) the presence (Figs 128 A – B) or the absence (Fig. 128 H) of radial depressions on the head valve; (2) the large variability of the height-width ratio in intermediate valves (Fig. 128 D, H / W = 0.39; Fig. 128 J, H / W = 0.43; Fig. 128 G, H / W = 0.60); (3) LA of intermediate valves scarcely differentiated from CA (Figs 128 C, 128 I) or raised and separated by a diagonal fold (Fig. 128 E). Comparisons. This species differs from the other Oligocene Lepidochitona because of the sculpture of CA and AMA (granules arranged in more or less distinct longitudinal striae), and LA and PMA (granules arranged in diverging striae running obliquely), vs. a sculpture with granules randomly arranged on the entire valve, typical of the species of Lepidochitona from Gaas (see Tabs 19, 21).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF194EE70FADFDA569C593A0.taxon	distribution	Distribution. Lower Oligocene: North Europe, Germany: Gienberg (Boettger 1869; Janssen 1978); Northeastern Atlantic: Paris Basin: France; Auvers-Saint-Georges, Morigny, Ormoy-La-Rivière (Janssen 1978; Lozouet & Maestrati 2012; this study); Belgium (Marquet et al. 2008).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF184EE90FADF9736F789458.taxon	description	Fig. 129	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF184EE90FADF9736F789458.taxon	materials_examined	Type material. The holotype (an intermediate valve) was lost due to war circumstances (Janssen 1978). Type locality. Gienberg (Germany). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene (Rupelian): Germany: Glimmerode: 7 valves (BD 954, Figs 129 A – B, 129 H), Gienberg: 2 valves (BD 955, Figs 129 C – D, 129 G). Upper Oligocene (Chattian): France: Abesse: 5 valves (BD 956, DA, MZB 50577, Figs 129 E – F); Ruisseau de l’église: 1 intermediate valve (JFL). Maximum width of the valves: 2.3 / 4 / 3.7 mm-	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF184EE90FADF9736F789458.taxon	description	Description. Head valve semicircular. Intermediate valves broadly rectangular (W / L = 2.41 – 3.10), semicarinate in anterior profile, moderately elevated (H / W = 0.39 – 0.40), anterior margin almost straight to slightly convex, side margins fairly straight, posterior margin almost straight, apex not very prominent, lateral area slightly raised, separated from central one by a distinct ridge. Tail valve semicircular, small, lightly projecting, subcentral mucro, antemucronal area separated from postmucronal one by a low ridge and a depression in front of that ridge. Tegmentum smooth and glossy, with prominent growth lines, aesthetes of same size. Articulamentum with apophyses wide, but not projecting far, insertion plates short, teeth irregular, slit formula 9 / 1 / 7, with a second slit very close to the posterior margin in intermediate valves, slit rays visible.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF184EE90FADF9736F789458.taxon	discussion	Remarks. This species was originally described by Sandberger (1859) based on a single intermediate valve from the lower Oligocene (Rupelian) of Germany. Unfortunately, the type material was lost during the Second World War. Later, Janssen (1978) described and illustrated valves from the type locality, as well as valves from late Oligocene localities in Germany, under the name Tonicella tenuissima, and attributed head and tail valves to this species. This species was placed in Tonicella Carpenter, 1873, by Janssen (1978), and later included by Sirenko & Dell’Angelo (2023) in the genus Lepidochitona Gray, 1821, widespread in the Tethys Ocean since the Eocene. Dell’Angelo et al. (2020 b) discussed a few intermediate and a single tail valve, from the Oligocene (Chattian) of the Aquitaine Basin, that agree with the features of L. tenuissima. The shape W / L of the intermediate valves of Lepidochitona tenuissima is variable, the valves are elongate, ca. 3 times as wide as long, W / L = 2.92 – 3.10, vs. 2.41 measured on the valve of L. tenuissima, figured by Janssen (1978: pl. 16, fig. 32); two valves from Waldbockelheim (Germany: BD collection) with a W / L ratio of 2.5 and 2.92, respectively, has been attributed to L. tenuissima. Comparisons. Lepidochitona tenuissima (Sandberger, 1859) differs from L. lira (Cherns & Schwabe, 2019) mainly by the smooth surface (vs. lateral areas rugose in L. lira), the posterior margin tapering to a pointed apex (vs. almost straight with apex not evident in L. lira) and the presence of a rib separating lateral and pleural areas (not present in L. lira). See Tab. 18 – 21 for comparison with the other Lepidochitona spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF184EE90FADF9736F789458.taxon	distribution	Distribution. Lower - upper Oligocene: North Europe, Germany: Böseberg, Gienberg, Glimmerode, Welschberg, Würzmühle (Sandberger 1859; Janssen 1978; Gürs 1983; this study); Upper Oligocene: northeastern Atlantic (Chattian): Aquitaine Basin, France: Abesse, Ruisseau de l’église (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EE90FADFE246B0196BC.taxon	type_taxon	Type species. Chiton ruber Linnaeus, 1767, by subsequent designation (Pilsbry 1893).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EE90FADFE246B0196BC.taxon	distribution	Distribution. Boreochiton is known from the Pliocene to the Recent, and presently distributed mainly in the northern Pacific, with one species in northern Atlantic (Sirenko 2016). The fossil record includes the Pleistocene of North Atlantic (Hoel 1914; Antevs 1917, 1928), Netherlands (Strack 2010) and Italy (Dell’Angelo & Giusti 1997).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EE90FADFE246B0196BC.taxon	discussion	Remarks. Sirenko (2000) reinstated the genus Boreochiton Sars, 1878, for some species originally attributed to the genus Tonicella, among which T. rubra (Linnaeus, 1767). The main feature of Boreochiton is the presence of a large fenestral gland in the tail of living individuals, only found in this genus. Boreochiton includes four species and one subspecies (Sirenko 2016; Sirenko & Dell’Angelo 2023). The main characters of Boreochiton ruber are included in Tab. 21.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	description	Fig. 130	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	materials_examined	Type material. The Linnean Society of London (Dodge 1952). Type locality. “ In Oceano Septentrionali instar Patellae affixa ”. Material examined. Pleistocene: Italy: Capraia Island-Capo Corso - 350 / 500 m: 327 valves (BD 957, Figs 130 A – K), Gallina: 3 valves (AV, BD 958, Fig. 130 L). Maximum width of the valves: 2.6 / 3.8 / 3.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	description	Description. Head valve semicircular, posterior margin widely V-shaped, some very faint radial fold may be present in some valves. Intermediate valves broadly rectangular (W / L = 2.40 – 2.79), moderately elevated (H / W = 0.35 – 0.53), semicarinate in anterior profile, anterior margin straight, side margins rounded, posterior margin straight to slightly concave at both sides of the protruding apex, lateral areas little or not elevated, hardly perceptible. Tail valve elliptical (W / L = 2.00 – 2.26), anterior margin roughly convex, straight between apophyses, mucro in anterior position, somewhat swollen, antemucronal and postmucronal slopes straight, generally rather steep. Tegmentum smooth to naked eye, sculptureless except for concentric growth lines, aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum strongly developed, apophyses wide, broadly triangular with rounded top, more or less trapezoid in the tail valve, separated by a rather narrow, flat jugal sinus, insertion plates short, slit formula 8 – 10 / 1 / 7 – 12, teeth sharp, smooth, slit rays little indicated, eaves finely porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	discussion	Remarks. This species has a long and rather complex nomenclatural history, summarized by Ferreira (1982) and Kaas & Van Belle (1985 b), triggered the differing opinions of Jakovleva (1952) and Sirenko (1974 a, 1974 b, 1976) on the identity of “ Tonicella ” rubra inhabiting the Pacific Ocean. Boreochiton ruber was seldom recorded from the Mediterranean (Leloup 1945; Fredj 1974; Monterosato 1890), unconfirmed by later authors (Dell’Angelo & Giusti 1997; Dell’Angelo & Smriglio 1999), so that is not included among the species living in the Mediterranean Sea (Renda et al. 2022; Dell’Angelo et al. 2024). As fossil, it has been reported from the Pleistocene (Sicilian) of Messina (Seguenza 1876) and Ficarazzi (Brugnone 1877; Tiberi 1877), but these records need to be confirmed. The species is ascertained from last glacial Pleistocene submerged assemblages between Capraia Island and Capo Corso (Ligurian Sea) at a depth of 350 / 500 m, where is common (Dell’Angelo & Giusti 1997; Dell’Angelo et al. 2024). The comparison between these fossil plates and recent material did not reveal any differences (compare figs 1 and 2 in Sirenko & Dell’Angelo 2023). The material examined shows a certain variability, mainly as regards the dorsal elevation (H / W = 0.35 – 0.53) and the greater or lesser accentuation of the apex of the intermediate valves, however always well evident. Comparisons. Boreochiton ruber (Linnaeus, 1767) is very similar to Tonicella marmorea (Fabricius, 1780); furthermore, their geographic and bathymetric ranges almost overlap. Loose valves of both genera are extremely difficult to be objectively assessed upon shell characters alone.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	distribution	Distribution. Pleistocene: North Atlantic: Russia, Barents Sea (Knipowitsch 1900), Spitsbergen (Hoel 1914; Hägg 1951), Norway (Brogger 1901), Sweden (Antevs 1917, 1928), Netherlands (Strack 2010); central Mediterranean, Italy: Capraia Island-Capo Corso - 350 / 500 m (Dell’Angelo & Giusti 1997; Dell’Angelo et al. 2001 a); Gallina (this study). Recent: N. Atlantic, Europe (Hansson 1998; Sneli & Gudmundsson 2018), Azores (Avila & Sigwart 2013). Genus Tonicella Carpenter, 1873	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	type_taxon	Type species. Chiton marmoreus Fabricius, 1780, by subsequent designation (Dall 1878).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	distribution	Distribution. Tonicella is known from the Miocene to the Recent. It occurs in the northern parts of the Pacific and Atlantic Oceans, and in the Arctic Ocean (Sirenko & Dell’Angelo 2023). Its fossil record extends back to the Miocene in Japan (Itoigawa et al. 1981 – 1982), Pliocene-Pleistocene of North America (Vendrasco et al. 2012 and references therein), Pleistocene of Europe (Feyling-Hanssen 1955) and California, U. S. A. (interglacial deposits, Muhs et al. 2006, 2012).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF164EEB0FADFC406F4D9604.taxon	discussion	Remarks. The valves in this genus are characterised by a largely smooth tegmental surface ornamented by tiny granules; the valves have weakly defined lateral areas (Ferreira 1982). The smooth ornament is an important feature of Tonicella, although also found in some species of Lepidochitona Gray, 1821. Tonicella likely originated in the North Pacific. Six species from the Eocene-Oligocene of Europe, before attributed to the genus Tonicella Carpenter, 1873, have been recently included in the genus Lepidochitona. The main characters of these species (with smooth tegmentum and pores of megalaesthetes and micraesthetes of the same size) are provided by Tab. 17 [for 5 species from the NE Atlantic area: L. adunca (Dell’Angelo, Lesport, Cluzaud & Sosso, 2020), L. lira (Cherns & Schwabe, 2019), L. modesta (Rolle, 1862), L. nuda (Dell’Angelo, Lesport, Cluzaud & Sosso, 2020) and L. redoniensis (Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018)] and in Tab. 19 [for the only species from the N Europe: L. tenuissima (Sandberger, 1859)]. The main characters of Tonicella marmorea (Fabricius, 1780) are defined in Tab. 21.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF144EEC0FADFC196E69906A.taxon	description	Fig. 131	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF144EEC0FADFC196E69906A.taxon	materials_examined	Type material. ZMK: Five well preserved specimens, dry, strongly curled, one selected as lectotype (the largest specimen, estimated length 4 cm) and the other four as paralectotypes by Ferreira (1982). Type locality. Greenland and Kragere (Norway) (Ferreira 1982: 117). Material examined. No fossil material available, only descriptions and illustrations from the literature. Recent: N. Atlantic: Europe: White Sea (BD 959, Figs 131 A – H), Barents Sea, Greenland, Iceland, Norway, Denmark; America: Canada (Nova Scotia), Maine (ca 25 spm). Maximum width: 4.2 / 4.5 / 3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF144EEC0FADFC196E69906A.taxon	description	Description. Head valve less than semicircular, posterior margin widely V-shaped, notched in the middle, slope slightly convex. Intermediate valves broadly rectangular (W / L = 2.73 – 2.75), moderately elevated (H / W = 0.37 – 0.44), semicarinate in anterior profile, anterior margin straight, except in valve ii where it is somewhat anteriorly produced, side margins more or less rounded, posterior margins about straight, apex decidedly perceptible, lateral areas hardly raised, separated from the central area by a shallow, diagonal depression, Tail valve small, semielliptical more than twice as broad as long (W / L = 2.43 – 2.71), very short, anterior margin evenly convex, posterior margin usually with a very shallow caudal sinus, mucro subcentral, not prominent, antemucronal slope almost straight, postmucronal slope about straight or slightly convex. Tegmentum uniformly, finely granulated in quincunx, growth lines relatively numerous, easily visible, concentrically crossing central areas, aesthetes very dense, each megalaesthete accompanied by many micraesthetes. Articulamentum with apophyses wide, short, anteriorly rounded, separated by a rather narrow, straight sinus, insertion plates short, slit formula 7 – 12 / l / 5 – 11, teeth slightly thickened at the edges, rugose on the outside, slit rays not indicated, eaves finely porous.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF144EEC0FADFC196E69906A.taxon	discussion	Remarks. Tonicella marmorea (Fabricius, 1780) has a long and rather complex nomenclatural history, summarized by Ferreira (1982) and Kaas & Van Belle (1985 b). Specimens from the East coast of the U. S. A. differ slightly from European specimens in being higher elevated, relatively narrower, their back more rounded (Kaas & Van Belle 1985 b). Our treatment of Tonicella marmorea is limited to the Pleistocene of the North Atlantic, based solely on citations of loose valves, without descriptions or figures, which could be questionable. Comparisons. Tonicella marmorea is very similar to Boreochiton ruber (Linnaeus, 1767).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF144EEC0FADFC196E69906A.taxon	distribution	Distribution. Pleistocene: North Europe: Russia: Barents Sea (Knipowitsch 1900), White Sea (Funder et al. 2002); Greenland: Harder et al. 1949; Norway (Brøgger 1901; Knipowitsch 1902; Holmboe 1904; Hoel 1907, 1914; Grønlie 1927; Hägg 1950, 1951; Feyling-Hanssen 1955); Sweden (Antevs 1917, 1928); Denmark (Petersen 2004). Recent: Arctic Ocean (Barents Sea, White Sea, The Kara Sea, near Spitzbergen and Franz Josef Land, and N. Canada); N. Atlantic Ocean, as far south as Massachusetts Bay on the western side, and the western coast of France on the eastern side; N. Pacific Ocean (Sea of Japan, the Okhotsk Sea and the Bering Sea, and near the Kurile, the Commander and the Aleutian Islands) (Ferreira 1982; Kaas & Van Belle 1985 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF134EEF0FADFA626E229604.taxon	type_taxon	Type species. Chiton fascicularis Linnaeus, 1767, by monotypy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF134EEF0FADFA626E229604.taxon	distribution	Distribution. Acanthochitona is known from the Oligocene to the Recent, the latter almost worldwide, except polar waters. The fossil record includes the Oligocene of New Zealand (Lee et al. 2014; Wu & Lee 2024), the Miocene of the Mediterranean Basin (Dell’Angelo et al. 2016, 2018 a, 2018 b), Paratethys (Šulc 1934; Bałuk 1984), West Pacific Islands (Ladd 1966), Australia (Cotton 1964), Pliocene to Pleistocene of the Mediterranean Basin (Dell’Angelo et al. 2001 a, 2013; Garilli et al. 2005), Australia (Cotton 1964) and New Zealand (Sutherland et al. 1995), Pleistocene of the Red Sea (Dell’Angelo et al. 2020 a), Holocene of Japan (Kuroda et al. 1980).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF134EEF0FADFA626E229604.taxon	discussion	Remarks. Acanthochitona is a confusing group; the standard chiton shell characters (e. g., shape of the valves, shape of granules and appearance of the jugal area) have a limited diagnostic value to easily set apart species (Ferreira 1983). Moreover, intraspecific variability within Acanthochitona species is high, rendering identification problematic (e. g., Leloup 1941, 1968). Six species of Acanthochitona are known as living along European waters [A. barbarae Dell’Angelo, Sosso & Taviani, 2024, A. crinita (Pennant, 1777), A. discrepans (Brown, 1827), A. fascicularis (Linnaeus, 1767), A. oblonga (Leloup, 1981) and A. pilosa Schmidt-Petersen, Schwabe & Haszprunar, 2015]; three of them are also known as fossil (A. crinita, A. fascicularis and A. oblonga) and considered in this study, whilst the occurrence of A. discrepans is doubtful (see appendix regarding the species of unclear taxonomic position). The main characteristcs of the species here discussed are reported in Tab. 22. Šulc (1934: p. 16) described the new subgenus Pseudoacanthochiton for species characterized by “ The larger number of cuts on the tail valve, of which two, however, that, based on their position, correspond to the incisions in the genus Acanthochiton, are much better developed than the other ones located in between them … This structure closely approaches [that of] the genus Acanthochiton, but the presence of a larger number of incisions forces us to place them in the genus Cryptoconchus ” (translation by A. Kroh). The subgenus Pseudoacanthochiton (Šulc, 1934) was subsequently synonymized with Acanthochitona Gray, 1821 (Smith 1960; Van Belle 1981; Bałuk 1984; Dell’Angelo et al. 2013), mainly highlighting that some tail valves of Acanthochitona with additional slits on the insertion plate had been reported [see above for A. fascicularis (Linnaeus, 1767)]. Anyway, all these reports illustrate valves that correspond in all respects to the valves of Acanthochitona, with the exception of the additional slits, which are, however, similar to the two slits typical of the genus Acanthochitona.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF104F100FADFC186A9A9343.taxon	description	Fig. 132	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF104F100FADFC186A9A9343.taxon	materials_examined	Type material. Holotype: MNHN. F. A 67148, tail valve, width 5.2 mm, Figs 132 E – F. Paratypes: MNHN. F. A 67149 – A 67154 (6 valves, Figs 132 B – C, 132 G), NHMW 2017 / 0108 / 0049 – 2017 / 0108 / 0054 (6 valves, Fig. 132 D), RGM. 1008416 – 1008421 (6 valves, Fig. 132 A). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Upper Miocene (Tortonian): Ligerian Basin, France: Beugnon: 6 valves (RGM. 1310173, RGM. 1310177), Renauleau: 53 valves (BD 156, MNHN. F. A 67156, NHMW 2017 / 0108 / 0056), Saint-Clément-de-la-Place: type material plus 1178 valves (BD 155, MNHN. F. A 67155, NHMW 2017 / 0108 / 0055, RGM. 1008358, RGM. 1008391, RGM. 1008422, RGM. 1008438). Lower Pliocene: Italy (Zanclean): Borzoli: 8 valves (BD 960, MSNG, Fig. 132 H). Upper Pliocene to upper Pleistocene: France: Bosq d’Aubigny: 7 valves (RGM. 1310181, RGM. 1310187). Maximum width of the valves: 5.3 / 7.8 / 7.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF104F100FADFC186A9A9343.taxon	description	Description. Head valve semicircular, posterior margin straight, apex hardly developed. Intermediate valve trapezoidal (WT / LT = 1.02 – 1.26), semicarinate in anterior profile, moderately elevated (H / W = 0.43), jugal margin straight or slightly convex, lateral margins straight, weakly rounded near posterior margin, posterior margin slightly concave either side of well-developed apex, jugal area triangular, smooth, raised, with presence of marked growth lines. Tail valve elliptical / polygonal, with greatest diameter along longitudinal axis (WT / LT = 0.80 – 0.86), mucro in posterior position, jugal area smooth, raised, antemucronal and postmucronal slopes almost straight, forming angle of about 50 ° with antemucronal slope. Tegmentum sculptured with irregularly arranged, ovate to elongate drop-shaped, flattened granules; several slightly more oblique longitudinal striae developed adjacent to JA. Granules widely separated from each other, of variable size, length 140 – 180 μm, slightly smaller on HV (ca. 110 – 120 μm). Each granule with one central megalaesthete and ca. 6 – 7 micraesthetes irregularly disposed. Articulamentum well developed, with large, quadrangular apophyses. The insertion plates are prominent — strongly protruding on the intermediate valves — and expanded to completely surround the tail valve, except along the jugal margin. Slit formula: 5 / 1 / 2.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF104F100FADFC186A9A9343.taxon	discussion	Remarks. The species is known only from the upper Miocene of France and Pliocene of Italy and France. Despite the large number of specimens found at Saint-Clément-de-la-Place, the valves are rarely complete and generally poorly preserved. The jugal area is smooth, except for the presence of marked growth lines. Some fine longitudinal furrows are sometimes visible when the surface is particularly eroded. The shape of the intermediate valves is variable, most of the valves show a length approximately equal to the width of the tegmental area (e. g., Fig. 132 B, WT / LT = 1.02) or slightly less, others show a much greater width than length (up to WT / LT = 1.26). Also, the shape of the tail valve is variable, tending to almost hexagonal in some valves. The micraesthetes are difficult to see, due to poor preservation of tegmentum. Comparisons. Acanthochitona andegavensis Dell’Angelo et al., 2018 is superficially similar to A. crinita (Pennant, 1777) (see below).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FF104F100FADFC186A9A9343.taxon	distribution	Distribution. Upper Miocene (Tortonian): northeastern Atlantic: France: Beugnon, Renauleau, Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b). Lower Pliocene: central Mediterranean: Italy: Borzoli (Dell’Angelo et al. 2021 b). Upper Pliocene-lower Pleistocene: northeastern Atlantic: France: Bosq d’Aubigny (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEF4F110FADF95D696592C7.taxon	description	Fig. 133	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEF4F110FADF95D696592C7.taxon	materials_examined	Type material. Holotype: MNHN. F. A 67157, tail valve, width 4 mm, Figs 133 F – H. Paratypes: MNHN. F. A 67158 – A 67163 (6 valves, Figs 133 A – C), NHMW 2017 / 0108 / 0057 – 2017 / 0108 / 0062 (6 valves, Figs 133 D – E), RGM. 1008423 – 1008425 (3 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Upper Miocene (Tortonian): France: Renauleau: 12 valves (BD 158, MNHN. F. A 67165), Saint-Clément-de-la-Place: type material plus 373 valves (BD 157, MNHN. F. A 67164, NHMW 2017 / 0108 / 0063, RGM. 1008359, RGM. 1008392, RGM. 1008426, RGM. 1008439), Sceaux d’Anjou: 4 valves (RGM. 1008445, RGM. 1008448). Maximum width of the valves: 5.3 / 6.8 / 5.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEF4F110FADF95D696592C7.taxon	description	Description. Head valve semicircular, posterior margin straight, apex scarcely evident. Intermediate valve trapezoidal (WT / LT = 1.19 – 1.34), semicarinate in anterior profile, rather elevated (H / W = 0.48), jugal margin straight to slightly convex, lateral margins straight, weakly rounded near posterior margin, posterior margin slightly concave on both sides of pronounced apex, jugal area triangular, smooth, raised, with the presence of prominent growth lines. Tail valve elliptical / polygonal, with greatest diameter along longitudinal axis (WT / LT = 0.84 – 0.92), mucro in slightly posterior position, jugal area smooth, raised, antemucronal and postmucronal slopes almost straight, forming angle of about 40 ° with antemucronal slope. Tegmentum sculptured with irregularly arranged, flattened roundish granules; granules densely packed, diameter 60 – 90 μm, with one central megalaesthete and up to 4 – 5 micraesthetes irregularly disposed. Articulamentum well developed, apophyses quadrangular, insertion plates well developed, strongly protruding on intermediate valves, expanded and completely surround tail valve, except for jugal margin, slit formula 5 / 1 / 2.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEF4F110FADF95D696592C7.taxon	discussion	Remarks. The studied material is poorly preserved, with valves abraded and seldom complete. JA is smooth and has prominent growth lines, some fine longitudinal furrows are sometimes visible when the surface is particularly eroded. The micraesthetes are difficult to see, due to poor preservation of tegmentum. Comparisons. At a first glance Acanthochitona chauvereauensis Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is closest to A. fascicularis (Linnaeus, 1767) in having the tegmentum sculpture formed by roundish and densely packed granules. It differs from A. fascicularis by a smooth JA (longitudinally striated in A. fascicularis), the different shape of the intermediate valves (more rectangular in A. fascicularis, with WT much greater than LT), the different shape of the tail valve (elliptical with WT> LT and with postmucronal slope strongly concave behind the mucro in A. fascicularis) and in lacking a deep incision in the granules, typical for A. fascicularis (see Fischer & Renner 1979: figs 4 – 5; Bonfitto et al. 2011: fig. 1).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEF4F110FADF95D696592C7.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Anjou, France: Renauleau, Saint-Clément-de-la-Place, Sceaux d’Anjou (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEED4F140FADFF056E7990FC.taxon	description	Fig. 134	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEED4F140FADFF056E7990FC.taxon	materials_examined	Type material. Neotype RSMNH 1978.052.02601, designated and figured by Kaas (1985 a: p. 591, fig. 27). Type locality. Hebrides Islands, Monach Island, North Uist, U. K. Material examined. Lower Miocene: France (Aquitanian): Lariey: 2 valves (JFL); France (Burdigalian): Coupe du fossé près de La Solitude: 1 valve (JFL), Gamachot: 58 valves (AC, BD 961, JFL, PR, MZB 50580 – 50582, Figs 134 B – D, 134 J – L), Maureilhan: 37 valves (BD 962, Fig. 134 A), Petit Bargues: 1 valve (JFL); Italy (Burdigalian): Sciolze: 5 valves (PG), Valle Ceppi: 1 valve (BD 963). Middle Miocene: France (Serravallian): Orthez: 1 valve (BD 964); Sallespisse Carré: 1 valve (DA); Italy (Langhian): Albugnano: 10 valves (BD 965, MZB 32108 – 32109, Figs 134 E – F, PG). Upper Miocene: Italy (Tortonian): Borelli: 2 valves (MGPT, MZB 32113), Montegibbio: 11 valves (BD 966, MZB 32112), Rio di Bocca d’Asino: 23 valves (BD 967, MZB 32110 - 32111, PG). Lower Pliocene: Italy: Borzoli: 67 valves (BD 968), Bussana: 252 valves (BD 969, Fig. 134 I, MZB 45767, Figs 134 G – H, SR), Caranchi: 8 valves (BD 970, MP), Garlenda: 6 valves (MP, MZB 45766), Genova Sestri: 22 valves (BD 971), Rio Sant’ Antonino: 105 valves (BD 972), Rio Torsero: 5 valves (BD 973, MZB 45768), Zinola: 1 valve (BD 974). Pliocene: Portugal: Vale de Freixo: 42 valves (BD 246, GeoFCUL VFX. 03.347, GeoFCUL VFX. 03.362 – 363, RGM. 1364020 – 1364021, MNHN. F. A 81992). Spain: El Papiol: 5 valves (BD 975); Estepona: 11 valves (BD 976); Italy: Piedmont: Baldichieri: 1 valve (BD 977), Montafia: 7 valves (BD 978), Vintebbio: 3 valves (BD 979); Emilia-Romagna: Cava di Campore: 20 valves (BD 980), Gagliardella “ Tagliata ”: 3 valves (BD 981); Tuscany: Bibbiano: 3 valves (BD 982), Castiglioncello del Trinoro: 1 valve (BD 983), Colle Val d’Elsa: 2 valves (BD 984), Montenero: 1 valve (BD 985), Orciano Pisano: 7 valves (BD 986), Pietrafitta: 15 valves (BD 987), Serre di Rapolano: 4 valves (BD 988); Latium: Magliano Sabina: 1 valve (BD 989); Sicily: Altavilla: 1 valve (BD 990), Trappeto: 13 valves (BD 991). Pleistocene: Italy: Emilia-Romagna: Torrente Stirone: 4 valves (BD 992); Tuscany: Capraia Island-Capo Corso - 350 / 500 m: 5 valves (BD 993), Cisternino: 2 valves (BD 994), Fauglia: 3 valves (BD 995), Riparbella: 25 valves (BD 996); Puglia: Gallipoli: 3 valves (BD 997); Calabria: Archi S. Francesco: 12 valves (BD 998), Carrabbati: 10 valves (BD 999), Gallina: 4 valves (BD 1000), Musalà: 1 valve (BD 1001), Pecoraro: 7 valves (BD 1002), Petti di Carrubbare: 2 valves (BD 1003), Pezzo: 4 valves (BD 1004), San Procopio: 3 valves (BD 1005), Stalettì: 1 valve (BD 1006) Vito Superiore: 1 valve (BD 1007); Sicily: Ficarazzi: 1 valve (BD 1008), Salice: 1 valve (BD 1009). Maximum width of the valves: 3.5 / 6 / 4.2 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEED4F140FADFF056E7990FC.taxon	description	Description. Head valve semicircular, posterior margin straight, slope slightly convex. Intermediate valves broadly rectangular (WT / LT = 1.43 – 1.70), rather elevated (H / W = 0.40 – 0.48), semicarinate in anterior profile, anterior margin straight between apophyses, lateral margins whose partly straight and partly rounded profile gives the valve a nearly pentagonal shape, posterior margin almost straight on both sides of pronounced apex, jugal area longitudinally striated, raised, not well separated from lateropleural area. Tail valve elliptical to circular (WT / LT = 1.16 – 1.35), jugal area longitudinally striated, raised, mucro well evident in subcentral to slightly posterior position, postmucronal area rather reduced, antemucronal slope slightly convex, postmucronal slope concave behind mucro. Whole tegmentum (except JA) sculptured with moderatly widespread, roundish / oval to drop or pear shaped granules, elevated, with flat or slightly concave surface, arranged along arched lines, jugal area longitudinally striated; granules of variable size, 1 – 2 times as long as wide, maximum size 194 μm, with a single posteriorly located megalaesthete and many micraesthetes, ranging from 12 to over 16, irregularly grouped mainly in anterocentral part of granules. Articulamentum well developed, apophyses quadrangular and strongly protruding, delimiting a wide jugal sinus, expanded and completely surrounding tail valve, except for jugal margin, slit formula 5 / 1 / 2.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEED4F140FADFF056E7990FC.taxon	discussion	Remarks. Acanthochitona crinita (Pennant, 1777) was reported by most Authors as A. fascicularis (non Linnaeus, 1767) before the work of Kaas (1985 a). Acanthochitona crinita is an extremely variable species with a complicated synonymy. The size and shape of the granules are highly variable, more or less drop-shaped or tending to an oval profile. Dell’Angelo et al. (2018 a) attributed tentatively to Acanthochitona crinita a single valve from the upper Oligocene (Chattian) of Saint-Etienne-d’Orthe (France, Aquitaine Basin). The scantness of material (1 valve) and geological antiquity makes highly unlikely its cospecificity with the extant taxon.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEED4F140FADFF056E7990FC.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Aquitanian-Burdigalian): Aquitaine Basin, France: Coupe du fossé près de La Solitude, Gamachot, Lariey, Maureilhan, Petit Bargues; Proto-Mediterranean Sea (Burdigalian): N. Italy: Sciolze, Valle Ceppi (Dell’Angelo et al. 2016); Middle Miocene: northeastern Atlantic (Serravallian): Aquitaine Basin, France: Orthez, Sallespisse Carré (Dell’Angelo et al. 2020 b); Proto-Mediterranean Sea (Langhian): Po Basin, N. Italy: Albugnano (Dell’Angelo et al. 2016); Central Paratethys (Langhian-Serravallian): Austria: Steinabrunn (Šulc 1934); Hungary: Devecser, Letkés (Dulai 2025 a, 2025 b); Poland: Korytnica, Rybnica (Bałuk, 1971, 1984; Studencka & Studencki 1988); Upper Miocene: Proto-Mediterranean Sea (Tortonian-Messinian): Po Basin, N. Italy: Borelli, Montegibbio, Rio di Bocca d’Asino (Dell’Angelo et al. 1999, 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: many localities (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013). Pliocene: northeastern Atlantic, Mondego Basin, Portugal (Dell’Angelo & Silva 2022); western Mediterranean, Estepona Basin, Spain (Dell’Angelo et al. 2004); central Mediterranean, Italy: many localities in Piedmont, Emilia-Romagna, Tuscany, Latium, Sicily (Dell’Angelo et al. 2001 a, 2013; Chirli 2004; Sosso & Dell’Angelo 2010; this study). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece (Koskeridou et al. 2009). Pleistocene: North Europe: Netherlands (Strack 2010); northeastern Atlantic: Portugal (Nobre 1892); central Mediterranean: Italy: many localities in Emilia-Romagna, Tuscany, Puglia, Calabria, Sicily (Dell’Angelo & Forli 1995 a; Dell’Angelo & Giusti 2000; Dell’Angelo et al. 2001 a; this study). Recent: Atlantic coast of Europe, from Scandinavia, N. Norway to 67.5 ° N (Hansson 1998), all the European coast (McKay & Smith 1979) to Spain and Portugal (Consolado Macedo & Borges 1999; Urgorri et al. 2017), Berlengas Arch. (Pisani Burnay 1986), plus Madeira (Kaas 1991), the Canary Islands (Kaas 1991; Hernández & Rolán 2011), and the Cape Verde Archipelago (Kaas 1991; Segers et al. 2009). Mediterranean Sea: Spain (Salas & Luque 1986), France, Italy (Dell’Angelo & Smriglio 1999); Croatia (Dell’Angelo & Zavodnik 2004); Greece and Aegean Sea Islands (Kattoulas et al. 1973; Strack 1988, 1990; Koukouras & Karachle 2005); Turkey (Ozturk et al. 2014); Israel (Barash & Danin 1977); Lebanon (Crocetta et al. 2014); Tunisia: Gulf of Gabes (Cecalupo et al. 2008); Black Sea (Mitov 2015).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEB4F180FADF98068EF94B4.taxon	description	Fig. 135	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEB4F180FADF98068EF94B4.taxon	description	Craspedochiton steinabrunensis; Studencka & Studencki 1988, tab. 2, 3.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEB4F180FADF98068EF94B4.taxon	materials_examined	Type material. Location currently unknown; not in NHMW. Type locality. Rudoltice (Czech Republik). Type stage. Middle Miocene. Material examined. Middle Miocene: Central Paratethys: Austria: Gainfarn: 2 valves (BD 1010, NHMW 1859 / 0027 / 0193), Niederleis: 2 valves (NHMW 2002 / 0087 / 0001); Czech Republic: Kninice: 1 valve (NHMW 1868 / 0001 / 0119), Lysice: 3 valves (NHMW 1865 / 0015 / 0162), Portzeich: 4 valves (NHMW 1871 / 0010 / 0211); Romania: Bujtur: 1 valve (NHMW 2010 / 0256 / 0022), Kostej: 10 valves (BD 1011, NHMW 2010 / 0256 / 0023, Figs 135 I – L), Lăpugiu de Sus: 20 valves (BD 1012, NHMW 1868 / 0001 / 0602, 2010 / 0256 / 0015, 2010 / 0256 / 0019, Figs 135 A – B); Hungary: Bánd: 33 valves (BD 1013), Borsodbóta: 1 valve (BD 1014), Letkés: 31 valves (BD 1015), Várpalota: 8 valves (BD 1016); Eastern Paratehys: Ukraine: Horodok: 42 valves (BD 1017), Kupin: 42 valves (BD 1018), Varovtsy: 500 + valves (BD 1019, Figs 135 C – H, 135 M – N), Velyka Levada: 30 valves (BD 1020), Zalistsi: 2 valves (BD 1021). Acanthochitona sp. I: Czech Republic: Hrusovany: NHMW 1861 / 0011 / 0202, intermediate valve (Figs 135 O – P, type figured by Šulc). Maximum width of the valves: 6 / 7 / 5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEB4F180FADF98068EF94B4.taxon	description	Description. Head valve semicircular, posterior margin straight, apex scarcely evident. Intermediate valves broadly rectangular (WT / LT = 1.45 – 1.89), moderately elevated (H / W = 0.37 – 0.64), semicarinate in anterior profile, posterior margin slightly concave on both sides of pronounced apex, jugal area generally narrow, longitudinally striated, raised, neatly separated from lateropleural area. Tail valve elliptical to circular, just straighter in jugal area (WT / LT = 1.20 – 1.70), jugal area longitudinally striated, raised, neatly separated from lateropleural area, mucro well evident in central position, antemucronal slope slightly convex, postmucronal slope concave behind the mucro. Whole tegmentum (except JA) sculptured with densely packed roundish to oval granules, rather elevated, irregularly arranged along arched lines, with flat or slightly concave surface, sometimes with a characteristic incision on surface, JA longitudinally striated; granules small, widely separated from each other, of variable size, maximum diameter 110 μm, with a central megalaesthete generally surrounded by 1 to 4 irregularly arranged micraesthetes. Articulamentum well developed, apophyses quadrangular and very protruding, delimiting a wide jugal sinus, expanded and completely surrounding tail valve, except for jugal margin, slit formula 5 / 1 / 2 +.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEB4F180FADF98068EF94B4.taxon	discussion	Remarks. Acanthochites faluniensis de Rochebrune, 1882 was described with the following diagnosis: “ Chiton (Acanthochites) fascicularis Lin. var. Reuss, loc. cit., p. 260, n ° 110, Tag. VIII, fig. 4, 6. A. --- Testa elongata, augusta; valvis intermediis, summitate sulcatis, sulcis latis profundis, utrinque longitudinaliter granatis; granis secessis; valva postica, lata, rotundata, intense rostrata, antice radiatim, postice concentrice granosa. Long. 0,002. Lat. 0,001. Hab. Sables de Rudelsdorf ”. And de Rochebrune adds: “ Reuss ne voit aucune différence essentielle, entre cette espèce et l’A. fascicularis Lin. et il la réunit provisoirement à elle … Une grande confusion règne danse le détermination des Acanthochites, compris dans le groupe du fascicularis de Linné ”. Considered conspecific with A. fascicularis (Linnaeus, 1767) by some Authors (e. g., Laghi 1977; Dell’Angelo et al. 1999, 2004, 2007 a), A. faluniensis differs from the extant species on some morphological characters (Bałuk 1971, 1984; Dulai 2001, 2005; Kroh 2003; Zágoršek 2006; Ruman & Hudácková 2015). The noteworthy variability of outlines of intermediate and tail valves, and of postmucronal slopes of tail valves of A. fascicularis is documented by Leloup (1941: figs 2 – 3; 1968: figs 4 – 6) and Dell’Angelo & Smriglio (1999: fig. 9) for Recent Mediterranean specimens. The putative differences between the two taxa are summarized in Studencka & Dulai (2010): • the tegmentum outline (the lateral margin of the tegmentum which usually is arched, rarely falciform in the posterior part in A. faluniensis, compared to more or less concave near the jugal area in A. fascicularis); • the ornamentation, with small, rounded and very crowded granules arranged in curved series in two directions, parallel to the jugum and radiating from it towards the outer margin in A. fascicularis; compared with finer granulation consisting of smaller numbers of distinctly larger rounded granules, each with 1 – 4 micraesthetes, arranged along orderly arched lines in A. faluniensis. The curved series of granules in in two directions in A. fascicularis is taken verbatim from the description of neotype of A. fascicularis designated by Kaas (1985 a: 585, figs 1 – 6); • the “ gentle ” postmucronal slope of the almost circular to more ellipsoidal tail valve of A. faluniensis, compared to deeply concave directly behind the sharp mucro in A. fascicularis. In our opinion, it is somehow subjective to identify the arrangement of granules in curved series radiating from the jugum as postulated; instead, while one or more series of granules may appear parallel to the jugum, other granules are disposed without any apparent order. Furthermore, SEM images do not seem to clearly document any distinct arrangement in striae of the granules in A. fascicularis (Fischer & Renner 1979). About the “ finer granulations ” of A. faluniensis, Fischer & Renner (1979) reported a granules’ width of 50 – 80 μm, with 0 – 3 micraesthetes for each central megalaesthete. Thus, a convincing separation between A. faluniensis and A. fascicularis based upon unquestionable morphological characters of loose valves seems extremely difficult at present. We prefer, however, to keep them provisionally distinct mainly because of their different paleobiogeographic domains.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEB4F180FADF98068EF94B4.taxon	description	As discussed above, the morphology of valves of Acanthochitona fascicularis and A. faluniensis is rather close, with respect to the shape of valves and granules. The shape of the valves of A. faluniensis shows a great variability, e. g., the head valve, more frequently with apex scarcely evident, but that can sometimes be very pronounced (see Ruman & Hudácková 2015: fig. 4.9). The shape of granules is variable, sometimes the surface shows a characteristic incision on the surface, as well figured by Fisher & Renner (1979: fig. 4 – 6) and Bonfitto et al. (2011: fig. 1) on living specimens of A. fascicularis. In some cases, the granules show a narrow ridge pointing roughly towards the apex of the valves (Figs 135 D, 135 P). We prefer at present to keep the two species distinct, pending future confirmation or otherwise of their possible synonymy, considering A. faluniensis a species restricted to the Paratethys. Comparisons. Acanthochitona faluniensis is morphologically close to A. fascicularis,	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEEB4F180FADF98068EF94B4.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria: Gainfarn, Niederleis, Steinabrunn (Šulc 1934; Kroh 2003; this study), Czech Republic: Drnovice, Knínice, Lysice, Rudoltice, Portzeich, Sudice, Vranovice, Židlochovice (Šulc 1934; Kroh 2003; Zágoršek, 2006; this study), Slovakia: Kúty, Rohožník, Devínska Nová Ves (Tomašových 1998; Kováč et al. 1999; Ruman & Hudácková 2015), Hungary: Bánd, Borsodbóta, Letkés, Szokolya, Várpalota (Studencka & Studencki, 1988; Dulai 2001, 2005; Dulai & Katona 2024; this study), Poland: Korytnica, Lychów, Miasteczko, Monastyrz, Navodzice, Rybnica, Trzesiny, Węglini, Weglinek (Bałuk 1965, 1971, 1984; Jakubowski & Musiał, 1977, 1979; Studencka & Studencki, 1988; Macioszczyk, 1988), Romania: Bujtur, Lăpugiu de Sus (Zilch 1934; Dell’Angelo et al. 2007 a; this study), Bulgaria: Trnienie (Jakubowski & Musiał, 1979); Eastern Paratehys: Ukraine: Horodok, Kupin, Olesko, Szuszkowce, Varovtsi, Velyka Levada, Zalistsi (Studencka & Dulai 2010; this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE74F1B0FADFE48683E9140.taxon	description	Fig. 136	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE74F1B0FADFE48683E9140.taxon	materials_examined	Type material. MNHN, Neotype designated and figured by Kaas (1985 a: p. 588, fig. 1). Type locality. “ in Barbaria ”; Neotype: Oran, Algeria. Material examined. Lower Miocene: France (Aquitanian): Lariey; 1 valve (JFL); France (Burdigalian): Carrière Vives: 1 valve (BD 1022), Gamachot: 3 valves (AC, BD 1023, MZB 50578, Figs 136 A – B), Le Thil: 1 valve (PR), Maureilhan: 4 valves (BD 1024); Italy (Burdigalian): Valle Ceppi: 2 valves (BD 1025). Middle Miocene: France (Serravallian): Carré: 1 valve (MZB 50579, Figs 136 C – D). Upper Miocene: France (Messinian?): Moulin-Pochas: 6 valves (PR); Italy (Tortonian): Borelli: 17 valves (BD 1026, MGPT, MZB 32106, PG), Montegibbio: 94 valves (BD 1027), Rio di Bocca d’Asino: 396 valves (BD 1028, MZB 32100 – 32105, Figs 136 I – L, PG), S. Agata Fossili: 10 valves (PG), Vigoleno: 2 valves (BD 1029), Villa Monti: 32 valves (BD 1030, PG). Lower Pliocene: Italy: Liguria: Borzoli: 366 valves (BD 1031), Bussana: 114 valves (BD 1032), Caranchi: 42 valves (BD 1033), Garlenda: 10 valves (MP), Genova Sestri: 5 valves (BD 1034), Rio Sant’ Antonino: 427 valves (BD 1035, Figs 136 E – F, MP, MZB 45761 – 45765), Rio Torsero: 36 valves (BD 1036, MP, PG, SR), Salea: 1 valve (BD 1037), Zinola: 2 valves (BD 1038). Pliocene: Spain: Barcelona, El Papiol: 2 valves (BD 1039), Estepona: 40 valves (BD 1040); Italy: Piedmont: San Damiano: 2 valves (VB), Vintebbio: 13 valves (BD 1041); Tuscany: Bibbiano: 11 valves (BD 1042), Castell’Anselmo: 3 valves (BD 1043), Castiglioncello del Trinoro: 7 valves (BD 1044), Cetona: 1 valve (BD 1045), Colle Val d’Elsa: 1 valve (BD 1046), Monte Antico: 1 valve (BD 1047), Montenero: 3 valves (BD 1048), Orciano Pisano: 75 valves (BD 1049), Pietrafitta: 62 valves (BD 1050), Pietrafitta Melograni: 32 valves (BD 1051), Pontedera: 2 valves (BD 1052), San Miniato: 9 valves (BD 1053), Serre di Rapolano: 8 valves (BD 1054); Emilia-Romagna: Castell’Arquato: 1 valve (BD 1055), Cava di Campore: 19 valves (BD 1056), Gagliardella: 6 valves (BD 1057); Umbria: Orvieto: 2 valves (BD 1058); Latium: Magliano Sabina: 1 valve (BD 1059), Monte Mario: 2 valves (BD 1060), Montelibretti: 1 valve (BD 1061); Sicily: Altavilla: 27 valves (AG, AR, BD 1062), Sciacca: 4 valves (BD 1063), Trappeto: 3 valves (BD 1064). Pleistocene: Italy: Tuscany: Caletta: 3 valves (BD 1065), Capraia Island-Capo Corso: 157 valves (BD 1066), Cisternino: 48 valves (BD 1067), Fauglia: 48 valves (BD 1068), Riparbella: 29 valves (BD 1069); Emilia-Romagna: Torrente Arda: 2 valves (BD 1070), Torrente Stirone: 12 valves (BD 1071); Puglia: Cutrofiano: 2 valves (BD 1072), Gallipoli: 4 valves (BD 1073), Marina di Novaglie: 2 valves (BD 1074), Porto Pirrone: 3 valves (BD 1075); Calabria: Archi S. Francesco: 81 valves (BD 1076), Bovetto: 1 valve (BD 1077), Carrabbati: 33 valves (BD 1078), Castellace: 2 valves (BD 1079), Gallina: 7 valves (BD 1080), Le Castella: 76 valves (BD 1081), Monasterace: 6 valves (BD 1082), Musalà: 4 valves (BD 1083), Pecoraro: 14 valves (BD 1084), Petti di Carrubbare: 20 valves (BD 1085), Pezzo: 234 valves (BD 1086), S. Maria di Catanzaro: 2 valves (BD 1087), San Procopio: 6 valves (BD 1088), Saracinello: 1 valve (BD 1089), Stalettì: 6 valves (BD 1090), Terreti: 24 valves (BD 1091), Torrente Boscaino: 3 valves (BD 1092), Vito Superiore: 1 valve (BD 1093); Sicily: Calderà: 2 valves (BD 1094), Capo Milazzo: 15 valves (BD 1095), Dattilo: 3 valves (BD 1096), Grammichele: 1 valve (BD 1097), Melilli: 1 valve (BD 1098), Menfi: 9 valves (BD 1099), Messina: 2 valves (BD 1100), Ogliastro: 2 valves (BD 1101), Salice: 2 valves (BD 1102), Selinunte Casa Catarinicchia: 3 valves (BD 1103), Selinunte Casa Parrino: 10 valves (BD 1104), Vittoria Case Buffa: 1 valve (BD 1105). Greece: Kyllini: 58 valves (BD 1106, DGUP, Figs 136 G – H), Perachora: 31 valves (BD 1107), Rhodes: Kritika: 6 valves (BD 1108). Maximum width of the valves: 5.3 / 8 / 7.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE74F1B0FADFE48683E9140.taxon	description	Description. Head valve semicircular, posterior margin straight, apex scarcely evident. Intermediate valves broadly rectangular (WT / LT = 1.60 – 2.10), moderately elevated (H / W = 0.37 – 0.48), semicarinate in anterior profile, posterior margin slightly concave on both sides of pronounced apex, jugal area generally narrow, longitudinally striated, raised, neatly separated from lateropleural area. Tail valve broadly circular, just straighter in jugal area (WT / LT = 1.20 – 1.29), jugal area longitudinally striated, raised, neatly separated from lateropleural area, mucro well evident in central position, antemucronal slope slightly convex, postmucronal slope strongly concave behind the mucro. Whole tegmentum (except JA) sculptured with densely packed roundish to oval granules, rather elevated, arranged along arched lines, with flat or slightly concave surface, sometimes with a characteristic incision in the middle of the granule posterior margin, JA longitudinally striated; granules small, widely separated from each other, of variable size, maximum diameter 93 μm, with a single central megalaesthete generally surrounded by 1 to 5 irregularly arranged micraesthetes (sometimes 0). Articulamentum well developed, apophyses quadrangular and very protruding, delimiting a wide jugal sinus, expanded and completely surrounding tail valve, except for jugal margin, slit formula 5 / 1 / 2 +.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE74F1B0FADFE48683E9140.taxon	discussion	Remarks. Acanthochitona fascicularis (Linnaeus, 1767) is an extremely variable species with a complicated synonymy. Kaas (1985 a) gives an accurate analysis of the taxonomic history of the three European species of Acanthochitona considered valid at the time, i. e. A. fascicularis, A. crinita (Pennant, 1777) and A. discrepans (Brown, 1827), Only Acanthochitona fascicularis and A. crinita occur in the Mediterranean Sea, while A. discrepans is known from northeastern Ireland and England. Before the work of Kaas (1985 a), Acanthochitona fascicularis was reported in the literature mainly as A. communis Risso, 1826, or A. discrepans (non Brown, 1827), while A. crinita was reported mainly as A. fascicularis (non Linnaeus, 1767). All Mediterranean fossil records of A. discrepans are here attributed to A. fascicularis; reports of A. discrepans from the English Pliocene (Coralline Crag, Sutton), could be related to the true A. discrepans. Laghi (1977) described the new species Craspedochiton (Pseudoacanthochitona) ambiguus upon valves from the lower Pliocene of “ La Tagliata ”, a locality near Modena. This species differs from A. fascicularis by the elliptical section granules (vs. those more rounded of A. fascicularis) and mainly by the tail valve with numerous slits (vs. 2 slits of A. fascicularis). Bałuk (1984) and Dell’Angelo et al. (1999) have already observed that the greatest number of slits in the tail valve may fall in the intraspecific variability of A. fascicularis, and therefore we agree with these authors in considering Craspedochiton ambiguus a junior synonym of A. fascicularis. Several tail valves with more than two slits have already been reported in the literature (e. g., Dell’Angelo et al. 2013: pl. 10, figs F – G; Dell’Angelo et al. 2016: pl. 5, figs 6 – 7, and this study: Figs 136 J – L). The great variability of the valves has been widely discussed (e. g., Leloup 1941, 1969; Dell’Angelo & Smriglio 1999; Dell’Angelo et al. 2013, 2016, 2020 b). Also, the shape of granules is variable, sometimes the surface shows a characteristic incision on the surface, as well figured by Fisher & Renner (1979: figs 4 – 6), Bonfitto et al. (2011: fig. 1) and Vončina et al. (2023: figs 4 C, 8 B) on living specimens. In the fossil material examined, are both valves with granules without (Fig. 136 F) and with incisions (Fig. 136 H), sometimes not as evident as those depicted for living specimens. Comparisons. Acanthochitona fascicularis is closest to A. faluniensis (de Rochebrune, 1882), from which it is practically indistinguishable (see above).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE74F1B0FADFE48683E9140.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Aquitanian-Burdigalian): Aquitaine Basin, France: Carrière Vives, Gamachot, Lariey, Le Thil, Maureilhan (Dell’Angelo et al. 2020 b); Proto-Mediterranean Sea (Burdigalian): N. Italy: Valle Ceppi (Dell’Angelo et al. 2016); Middle Miocene: northeastern Atlantic (Serravallian): Aquitaine Basin, France: Sallespisse (Dell’Angelo et al. 2020 b); Upper Miocene: northeastern Atlantic: Ligerian Basin, France: Moulin Pochas (Dell’Angelo et al. 2020 b); Proto-Mediterranean Sea (Tortonian): Po Basin, N. Italy: Borelli, Montegibbio, Rio di Bocca d’Asino, S. Agata Fossili, Vigoleno, Villa Monti (Laghi 1977; Dell’Angelo et al. 1999, 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: many localities (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b). Pliocene: western Mediterranean, Estepona Basin, Spain (Mancini 1998; Dell’Angelo et al. 2004, 2012); central Mediterranean, Italy: many localities in Piedmont, Emilia-Romagna, Sicily, Umbria, Latium, Sicily (Malatesta 1962; Laghi 1977; Dell’Angelo et al. 2001 a; Chirli 2004; this study). Upper Pliocene to upper Pleistocene: central Mediterranean, Greece (Koskeridou et al. 2009). Pleistocene: North Atlantic: Netherlands (Strack 2010); central Mediterranean, Italy: many localities in Emilia-Romagna, Tuscany, Puglia, Calabria, Sicily (Malatesta 1962; Dell’Angelo & Forli 1995 a; Dell’Angelo et al. 2001 a; this study), Greece: Kyllini, Perachora (Garilli et al. 2005; this study), Tunisia: Monastir (Castany et al. 1956). Recent: A tlantic coast of Europe: from the English Channel and Bretagne (McKay & Smith 1979) to Spain and Portugal (Borja 1987; Rolan Mosquera et al. 1990; Consolado Macedo et al. 1999; Strack 2010; Urgorri et al. 2017), Berlengas Arch. (Pisani Burnay 1986), plus the Azores (Kaas 1991; Avila & Sigwart 2013), Madeira (Kaas 1991) and Selvagens Arch. (Kaas 1991) and the Canary Islands (Kaas 1991; Hernández & Rolán 2011). Mediterranean Sea: Spain (Salas & Luque 1986), France, Italy (Dell’Angelo & Smriglio, 1999); Croatia (Dell’Angelo & Zavodnik 2004); Greece and Aegean Sea Islands (Kattoulas et al. 1973; Strack 1988, 1990; Koukouras & Karachle 2005); Turkey (Ozturk et al. 2014); Israel (Barash & Danin 1977); Lebanon (Crocetta et al. 2014); Tunisia (Kaas 1989; Cecalupo et al. 2008); Black and Marmara Sea (Öztürk et al. 2014).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE44F1C0FADFB5C683A9378.taxon	description	Fig. 137	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE44F1C0FADFB5C683A9378.taxon	materials_examined	Type material. Holotype: RGM. 1008383, tail valve, width 3.3 mm, Figs 137 F – H. Paratypes: MNHN. F. A 67139 – A 67144 (6 valves), Figs 137 A – D, NHMW 2017 / 0108 / 0042 – 2017 / 0108 / 0047 (6 valves), RGM. 1008384 – 1008389 (6 valves). Type locality. Saint-Clément-de-la-Place (France). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): France: Renauleau: 12 valves (BD 154, MNHN. F. A 67147,), Saint-Clément-de-la-Place: type material plus 884 valves (BD 153, MNHN. F. A 67145 – A 67146, Fig. 137 E, NHMW 2017 / 0108 / 0048, RGM. 1008357, RGM. 1008390, RGM. 1008415, RGM. 1008437). Maximum width of the valves: 2.5 / 4.8 / 3.3 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE44F1C0FADFB5C683A9378.taxon	description	Description. Head valve semicircular, posterior margin straight, apex scarcely developed. Intermediate valve broadly rectangular, rounded in anterior profile, also longitudinally, rather elevated (H / W = 0.45), anterior margin almost straight or weakly convex, posterior margin slightly concave either side of strongly protruding apex, jugal area triangular, longitudinally striated, not raised. Tail valve elliptical, mucro slightly posterior, jugal area longitudinally striated, not raised, antemucronal and postmucronal slopes almost straight, forming an angle of about 55 – 60 ° with antemucronal slope. Tegmentum sculptured with elongated flattened granules, irregularly arranged in radial striae starting from apex (on HV and intermediate valves) or from mucro on tail valve. Granules widely separated, of variable size, up to 250 μm in length on intermediate valves, slightly smaller on tail valve, with one central megalaesthete and many micraesthetes, 15 or more. Articulamentum poorly preserved, apophyses quadrangular, narrow, insertion plates extending far over the tegmentum, slit formula 5 / 1 / 2.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE44F1C0FADFB5C683A9378.taxon	discussion	Remarks. The species is known from the Miocene (Tortonian) of France (Dell’Angelo et al. 2018 b). We analysed a large collection of valves, rarely complete and poorly preserved. The sculpture of the intermediate valves of this species is highly characteristic, with the radial striae of elongated flattened granules radiating from the apex, which continue from longitudinal striae on the jugal area without interruption, the jugal area is not raised and the profile of the valve is evenly rounded. The micraesthetes are difficult to see because of the poor preservation of tegmentum, although 15 at least are visible on granules in one case (Fig. 137 D). Comparisons. Acanthochitona globosa Dell’Angelo, Landau, Van Dingenen & Ceulemans, 2018 is superficially similar to A. oblonga (Leloup, 1981) in having the tegmentum sculpture formed by elongated granules. Acanthochitona globosa differs from A. oblonga by the different shape of the intermediate and tail valves (profile semicarinate and with JA raised and distinctly separated from the other areas in A. oblonga), the less developed articulamentum and the greater number of the micraesthetes in the pustules (15 or more in A. globosa vs. 6 – 9 in A. oblonga).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE44F1C0FADFB5C683A9378.taxon	distribution	Distribution. Upper Miocene: northeastern Atlantic (Tortonian): Anjou, France: Renauleau, Saint-Clément-de-la-Place (Dell’Angelo et al. 2018 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	description	Fig. 138	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	materials_examined	Type material. Holotype: MZB 32107, tail valve, width 7.5 mm (Figs 138 A – D). Type locality. Borelli (Piedmont, Italy). Type stage. Upper Miocene (Tortonian).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	etymology	Etymology. We name this species after Mathias Harzhauser in recognition of his outstanding contribution to the taxonomy and biogeography of European Cenozoic Mollusca.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	materials_examined	Material examined. Upper Miocene: Italy: Borelli: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	diagnosis	Diagnosis. Tail valve elliptical / polygonal, jugal area neatly separated from lateropleural area, mucro well evident in posterior position. Whole tegmentum (except JA) sculptured with densely packed, elevated, irregularly arranged roundish granules. Articulamentum particularly expanded, more than three times width of tegmentum area, apophyses very protruding.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	description	Description. Head and intermediate valves unknown. Tail valve elliptical / polygonal, with greatest diameter along longitudinal axis (WT / LT = 0.81), jugal area neatly separated from lateropleural area, mucro well evident in posterior position, antemucronal and postmucronal slopes almost straight. Whole tegmentum (except JA) sculptured with densely packed roundish granules, flattened at top, elevated, irregularly arranged. Articulamentum particularly expanded, more than three times width of tegmentum area, apophyses very protruding, completely surrounding tail valve, except for wide jugal margin, two slits.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	discussion	Remarks. The only valve available is strongly characterised by the reduced tegmentum and the strongly protruding apophyses, a feature not present in any other European Acanthochitona spp. The jugal area is eroded, so that its sculpture cannot be appreciated. Comparisons. The species shares the large expansion of the articulamentum with Acanthochitona mastalleri Strack, 1989, an extant species from the Red Sea, also recorded from the late Pleistocene (Dell’Angelo et al. 2020 a), which differs in having such diminished tegmentum that the anterior part of the tail valve is only the jugum without lateropleural areas with granule sculpture.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE34F1D0FADF9056FFB900C.taxon	distribution	Distribution. Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N. Italy: Borelli (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE24F1E0FADFA106846929A.taxon	description	Fig. 139	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE24F1E0FADFA106846929A.taxon	materials_examined	Type material. Holotype MU, one specimen, 14 mm long. Type locality. Salina Bay (Malta), “ under rocks at 3 meters depth ”. Material examined. Upper Miocene: Italy: Montegibbio: 3 valves (BD 1109, MZB 32114, Figs 139 A – B), Rio di Bocca d’Asino: 1 valve (BD 1110). Lower Pliocene: Italy: Liguria: Bussana: 7 valves (BD 1111, MZB 45769 – 45770, Figs 139 C – D), Rio Sant’ Antonino: 1 valve (MP). Pleistocene: Italy: Tuscany: Riparbella: 8 valves (BD 1112). Recent: S. Foca (Italy): valves (BD 3811, Figs 139 E – H). Maximum width of the intermediate valves: 4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE24F1E0FADFA106846929A.taxon	description	Description. Head valve semicircular, posterior margin straight, slope slightly convex. Intermediate valves broadly rectangular (WT / LT = 1.73), semicarinate in anterior profile, moderately elevated (H / W = 0.42), anterior margin straight between apophyses, posterior margin almost straight on both sides of pronounced apex, jugal area longitudinally striated, raised. Tail valve broadly circular (WT / LT = 1.29), jugal area longitudinally striated, raised, mucro well evident, subcentral, antemucronal slope almost straight, postmucronal slope slightly concave. Whole tegmentum (except jugal area) sculptured with widely spread, oval, strongly elongated granules, irregularly arranged in radial striae, JA longitudinally striated; granules of variable size, 3 - 5 times as long as wide, maximum size 275 μm, with a single posteriorly located megalaesthete generally surrounded by 6 to 9 micraesthetes, irregularly grouped mainly in middle part of granules. Articulamentum well developed, apophyses quadrangular and strongly protruding, delimiting a wide jugal sinus, slit formula - / 1 / -.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE24F1E0FADFA106846929A.taxon	discussion	Remarks. This species, described by Leloup based on 4 specimens collected at Salina Bay (Malta), is characterized by the very extended sharp granules of the tegmentum. After the original description, it has been reported from other Mediterranean localities (Dell’Angelo & Smriglio 1999); large series from the Apulian Adriatic coast demonstrate greater variability of granules with respect to Leloup’s original description. Acanthochitona oblonga (Leloup, 1981) is so similar to A. crinita (Pennant, 1777) that subsequent authors (e. g., Kaas 1985 a; Dell’Angelo & Smriglio 1999) consider them as conspecific, until Bonfitto et al. (2011) demonstrated its specific validity based on morphological and molecular argunments. In paleontological legacy the species is known as intermediate valves from the Middle Miocene of Paratethys, upper Miocene of N. Italy, lower Pliocene of Liguria and the Pleistocene of Riparbella. Comparisons. See Tab. 22 for a comparison with the Acanthochitona spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE24F1E0FADFA106846929A.taxon	distribution	Distribution: Middle Miocene: Central Paratethys (Langhian-Serravallian): Hungary: Várpalota (Dulai & Katona 2024). Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N. Italy: Montegibbio, Rio di Bocca d’Asino (Dell’Angelo et al. 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: Bussana, Rio Sant’Antonino (Dell’Angelo et al. 2013). Pleistocene: central Mediterranean, Italy: Riparbella (Dell’Angelo & Forli 1995 a). Recent: Southern Mediterranean Sea in scattered Italian localities, Malta, Cyprus, Tunisia (Dell’Angelo & Smriglio 1999; Bonfitto et al. 2011).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE04F000FADFF056BA79490.taxon	description	Fig. 140	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE04F000FADFF056BA79490.taxon	materials_examined	Type material. Holotype NHMW 2010 / 0256 / 0001, intermediate valve, width 4.18 mm, Figs 140 A – E. Type locality. Pötzleinsdorf (Austria). Type stage. Middle Miocene. Material examined. Middle Miocene: Central Paratethys: Austria: Pötzleinsdorf: type material, plus 1 valve (NHMW 2010 / 0256 / 0008), width 3.2 mm, Figs 140 F – H; Hungary: Letkés: 2 valves (BD 1157), Figs 140 K – L; Romania: Lăpugiu de Sus: 2 valves (BD 1156), Figs 140 I – J. Maximum width: 7.4 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE04F000FADFF056BA79490.taxon	description	Description. Head and tail valves unknown. Intermediate valves broadly rectangular (W / L = 1.94), scarcely elevated (H / W = 0.29 – 0.36), semicarinate in anterior profile, posterior margin slightly concave on both sides of pronounced apex, jugal area longitudinally striated, not raised, neatly separated from lateropleural area. Whole tegmentum (except JA) sculptured with densely packed roundish granules, rather elevated, irregularly arranged along arched lines, with flat surface, JA longitudinally striated; granules small, widely separated from each other, roundish (diameter 46 – 57 µm), extended with a short longitudinal rib, more oval in LA (up to 65 µm), with a central megalaesthete generally surrounded by 2 – 3 irregularly arranged micraesthetes. Articulamentum well developed, apophyses quadrangular and very protruding, delimiting a wide jugal sinus, one slit in intermediate valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE04F000FADFF056BA79490.taxon	discussion	Remarks. Acanthochitona plana (Šulc, 1934) was described based on a single intermediate valve from Pötzleinsdorf (Austria), flatter and with an ornamentation identical with that of A. faluniensis (de Rochebrune, 1882). The material present at NHMW is represented by the holotype (the intermediate valve figured by Šulc, Figs 140 A – E) and two other valves (intermediate and tail) from the same type locality, Pötzleinsdorf, of which the tail valve is here attributed to A. sandeciana Bałuk, 1965 (see below). Comparisons. Acanthochitona plana is morphologically close to A. faluniensis, from which it differs mainly by the smaller granules and the rather flattened valves.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEE04F000FADFF056BA79490.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria: Pötzleinsdorf (Šulc 1934; Sieber 1953, 1959; this study); Hungary: Letkés (this study); Romania: Delineşti (Stancu & Andreescu 1968), Lăpugiu de Sus (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	description	Fig. 141	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	materials_examined	Type material. Syntypes in the Bałuk’s collection, three valves (head, intermediate and tail, figured in Bałuk (1965: pl. 1, figs 9 – 11), Figs 141 A – C. Type locality. Niskowa (Poland) Type stage. Middle Miocene. Material examined. Middle Miocene: Paratethys: Austria: Pötzleinsdorf: NHMW 2010 / 0256 / 0008, tail valve [determined as Acanthochitona planus (Šulc, 1934)], width 2 mm (Figs 141 F – H); Hungary: Bánd: 5 valves (BD 1158), Figs 141 D – E. Maximum width of the valves: 4.6 / 4.5 / 4 mm (from Bałuk 1965).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	description	Description. Head valve semicircular, posterior margin straight. Intermediate valves broadly rectangular, moderately elevated, posterior margin slightly concave on both sides of pronounced apex, jugal area longitudinally striated, raised, neatly separated from lateropleural area. Tail valve broadly circular, just straighter in jugal area, WT / LT = 1.22, jugal area longitudinally striated, raised, neatly separated from lateropleural area, mucro well evident in subcentral position, antemucronal slope straight, postmucronal slope slightly concave just behind mucro. Whole tegmentum (except JA) sculptured with densely packed roundish granules, flattened at top, elevated, irregularly arranged, JA longitudinally striated; granules small (diameter 50 – 55 µm), widely separated from each other, with a single megalaesthete generally surrounded by 1 to 3 irregularly arranged micraesthetes. Articulamentum well developed, apophyses quadrangular and very protruding, delimiting a wide jugal sinus, expanded and completely surrounding tail valve, except for jugal margin, slit formula 5 / 1 / 2.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	discussion	Remarks. Acanthochitona sandeciana Bałuk, 1965 has been described upon material from the Middle Miocene of Niskowa (Poland), and more recently from Korytnica (Poland) (Bałuk 1984) and Rohožník (Slovakia) (Ruman & Hudácková 2015). This species is characterized by the tail valve with an esagonal shape (as Acanthochiton sp. III Šulc, 1934 from Pötzleinsdorf, Austria), and the tegmentum sculpture like that of A. faluniensis, but with “ smaller and more densely packed granules ” (Bałuk 1984, p. 292). The tail valve from Pötzleinsdorf (Austria) at NHMW (Fig 141 E) has a shape which corresponds entirely to that described by Šulc (1934: text-fig. 4) as Acanthochiton sp. III, also in the form of apophyses, and comes from the same locality (Pötzleinsdorf) as the plate described by Šulc. We attribute also tentatively to Acanthochitona sandeciana some valves from Bánd, one of which figured (Figs 141 D – E). Comparisons. See Tab. 22 for a comparison with the Acanthochitona spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria: Pötzleinsdorf (Šulc 1934; this study), Poland: Korytnica, Niskowa (Bałuk 1965, 1984), Slovakia: Rohožník (Ruman & Hudácková 2015), Hungary: Bánd (this study). Genus Craspedochiton Shuttleworth, 1853	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	type_taxon	Type species. Chiton laqueatus Sowerby, 1842, by monotypy.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	distribution	Distribution. Craspedochiton is known from the Eocene to the Recent, with a living distribution in the Atlantic (African coast), and temperate and (sub-) tropical Indo-Pacific (Schwabe & Els 2019). The fossil record extends back to the upper Eocene or lower Oligocene in Washington, U. S. A. (Dell’Angelo et al. 2011), the Oligocene in France (Dell’Angelo et al. 2020 b), the Miocene-Pleistocene in Europe (Šulc 1934; Kroh 2003; Dell’Angelo et al. 2004, 2013, 2016, 2020 b; Garilli et al. 2005; Studencka & Dulai 2010), the Pliocene-Pleistocene of New Zealand (Sutherland et al. 1995), the Pleistocene deposits of the Red Sea (Dell’Angelo et al. 2020 a)	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFF4F010FADFE6C6F309016.taxon	discussion	Remarks. Eight species are currently attributed to Craspedochiton in the Oligocene - Pleistocene of Europe. Schwabe & Els (2019) presented an emended diagnosis of the genus Craspedochiton, agreeing with the features of the species of Craspedochiton discussed herein. The main characters of the species are reported in Tab. 23.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFE4F060FADFAEF694893A4.taxon	description	Fig. 142	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFE4F060FADFAEF694893A4.taxon	materials_examined	Type material. Neotype MZB 7062 (head valve), designated by Dell’Angelo & Palazzi (1989: fig. 1). Type locality. Altavilla Milicia (Sicily, Italy). Type stage. Pliocene-Pleistocene (upper Piacenzian to lower Gelasian) (Dominici et al. 2020). Material examined. Lower Miocene (Burdigalian): France: Carrière Vives: 10 valves (BD 1113, MZB 50584 – 50585), Pont St Martin: 1 valve (JFL); Italy: Valle Ceppi: 3 valves (MZB 32115 – 32116, PG). Middle Miocene: Central Paratethys: Austria: Forchtenau: 2 valves (NHMV 1866 / 0001 / 1209, as Cryptoconchus (Craspedochiton) profascicularis, Figs 143 M – O); Hungary: Letkés: 1 valve (BD 1114). Upper Miocene (Tortonian): Italy: Borelli: 8 valves (BD 1115, MZB 32119 – 32120, Fig. 142 C, MGPT, PG), Montegibbio: 2 valves (LB), Rio di Bocca d’Asino: 8 valves (BD 1116, MZB 32117 – 32118, PG). Lower Pliocene (Zanclean): Italy: Borzoli: 66 valves (BD 1117, Figs 142 H – I, MZB 45773), Caranchi: l valve (MP), Garlenda: 3 valves (MP), Rio Sant’ Antonino: 47 valves (BD 1118, MP, MZB 45772, PG), Rio Torsero: 2 valves (BD 1119, PG), Zinola: 1 valve (MZB 45771). Pliocene: Spain: Estepona: 17 valves (BD 1120); Italy: Piedmont: Valle Andona: 5 valves (BD 1121, NHMV 1877 A / 0018 / 0088, the tail valve described and figured by Šulc, as Cryptoconchus (Craspedoplax) sp. I, Fig. 143 P), Valle Botto: 1 valve (BD 1122); Tuscany: Bibbiano: 1 valve (BD 1123), Castell’Anselmo: 1 valve (BD 1124), Cigliano: 1 articulated spm (MZUB); Luciana: 2 valves (BD 1125), Orciano Pisano: 6 valves (BD 1126), Pietrafitta Melograni: 1 valve (BD 1127), Serre di Rapolano: 6 valves (BD 1128), Volterra: 1 valve (BD 1129); Emilia-Romagna: Bacedasco: 1 valve (BD 1130), Cava di Campore: 5 valves (BD 1131); Umbria: Cava di Ficulle: 1 valve (BD 1132), Monteleone d’Orvieto: 2 valves (BD 1133); Latium: Magliano Sabina: 2 valves (BD 1134); Sicily: Altavilla: 100 valves (AG, AR, BD 1135, Figs 142 D, 142 J – L); Sicily: Trappeto: 3 valves (BD 1136). Pleistocene: Italy: Sicily: Dattilo: 1 valve (BD 1137, Figs 142 E – G); Greece: Kyllini: 5 valves (BD 1138, DGUP, Figs 142 A – B). Maximum width of the valves: 9.8 / 27 / 8.6 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFE4F060FADFAEF694893A4.taxon	description	Description. Head valve semicircular, with five well evident radial ribs, slope almost straight. Intermediate valve polygonal (WT / LT = 1.78 – 2.75), semicarinate in anterior profile, moderately elevated (H / W = 0.23 – 0.42), anterior margin almost straight in jugum, side margins straight, posterior margin straight or slightly concave at both sides of prominent apex, jugal area large, triangular, lateral areas well separated from central area by a large diagonal fold. Tail valve elliptical (WT / LT = 1.62 – 1.89), anterior margin straight or slightly concave between apophyses, mucro subcentral, not elevated and poorly developed, antemucronal slope straight or slightly convex, postmucronal slope concave just behind mucro. Tegmentum covered with large, elevated granules of irregular shape, except on JA; shape, elevation and density of granules and granulation patterns highly variable, from single granules, regularly ellipsoidal, to coalescent granules forming irregular cords, JA smooth, top of granules convex. Each granule with many megalaesthetes and micraesthetes (up to 20 – 25 or more) arranged without any apparent order. Articulamentum with trapezoidal, large apophyses, expanded in tail valve, large posterior callus in tail valve, slit formula 5 / 1 / 7 – 8, insertion lamina reduced to a rather narrow band in head valve, teeth broad, roughed dorsally, more irregular in tail valves, slits deep, dorsally slightly elevated, forming channel.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFE4F060FADFAEF694893A4.taxon	discussion	Remarks. Craspedochiton altavillensis (Seguenza, 1876) is characterized by a tegmentum covered with large, elevated granules of irregular shape, except on JA (smooth). The synonymy of Craspedochiton deslongchampsi (de Rochebrune, 1882) with C. altavillensis was established and discussed by Dell’Angelo & Palazzi (1988). Dell’Angelo et al. (2003) reported a complete, articulated specimen of Craspedochiton altavillensis from the Italian Pliocene, the only case known to date.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFE4F060FADFAEF694893A4.taxon	description	Two valves (head and intermediate) from Forchtenau are present in the Šulc collection at NHMW. The head valve (Fig. 142 M) is identical to the holotype of Boettger figured by Zilch, and to the neotype of Craspedochiton altavillensis designated by Dell’Angelo & Palazzi (1988). The comparison of these two valves enabled Dell’Angelo et al. (1999, 2016) to consider Cryptoconchus (Craspedochiton) profascicularis conspecific with Craspedochiton altavillensis, and we confirm this synonymy. The incomplete intermediate valve (Figs 142 N – O) is consistent with the attribution to C. altavillensis, by its pentagonal layout, the raised rib separating the lateral and lateropleural areas, the tegmentum covered with large, elevated granules of irregular shape, except for JA. The tail valve figured by Šulc (1934: pl. 1, figs 20 – 21) as Cryptoconchus (Craspedoplax) sp. I, from the Pliocene of Valle Andona (Italy), is present in the Šulc collection at NHMW (Fig. 142 P). We agree that this valve is within the range of variability of Craspedochiton altavillensis, as already reported by Laghi (1977) and Dell’Angelo et al. (1999). Dell’Angelo et al. (2016) reported Craspedochiton altavillensis from the lower Miocene (Burdigalian) of the Turin Hills, Piedmont up to the Pleistocene of Greece and Italy. Such an extended stratigraphic range with different paleoclimatic scenarios and paleobiogeographic distribution is suspect, and may suggest that it may be a group of morphologically closely-related species. Dell’Angelo et al. (2016) have provisionally adopted a conservative approach and used the species name in a broad sense, pending further study. Comparisons. The species of Craspedochiton with the closest sculpture to C. altavillensis (large irregular granules tending to coalesce) is C. lozoueti Dell’Angelo, Lesport, Cluzaud & Sosso, 2020, which however differs from C. altavillensis (Seguenza, 1876) mainly for the greater elevation of the valves (more flattened in C. altavillensis), LA not separated from CA (well separated by a large diagonal fold in C. altavillensis), and the different shape of the tail valve. See Tab. 23 for a comparison with the Craspedochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEFE4F060FADFAEF694893A4.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Carrière Vives, Pont St Martin (Dell’Angelo et al. 2020 b); Proto-Mediterranean Sea (Burdigalian): N. Italy: Valle Ceppi (Dell’Angelo et al. 2016); Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria: Forchtenau (Šulc, 1934), Poland: Korytnica (Bałuk 1971, 1984), Hungary: Borsodbóta, Letkés (Schwabe & Dulai 2024; this study), Romania: Costej (Boettger 1907; Zilch 1934; Dell’Angelo et al. 1999); Eastern Paratehys: Ukraine: Varovtsi (Studencka & Dulai 2010); Upper Miocene: Proto-Mediterranean Sea (Tortonian): N. Italy: Borelli, Montegibbio, Rio di Bocca d’Asino (Laghi 1977; Dell’Angelo et al. 1999, 2016). Lower Pliocene: central Mediterranean, Italy: Liguria: many localities (Sosso & Dell’Angelo 2010; Dell’Angelo et al. 2013, 2021 b). Pliocene: western Mediterranean, Estepona Basin, Spain: Estepona (Dell’Angelo et al. 2004); central Mediterranean, Italy: many localities in Piedmont, Emilia-Romagna, Tuscany, Umbria, Latium, Sicily (Dell’Angelo et al. 2001 a, 2012, 2013; Chirli 2004; Sosso & Dell’Angelo 2010; this study). Pleistocene: central Mediterranean, S. Italy: Dattilo (Garilli 1998, 2011; this study), Greece: Kyllini (Garilli et al. 2005).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF94F070FADF9786B6B90C5.taxon	description	Fig. 143	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF94F070FADF9786B6B90C5.taxon	materials_examined	Type material. Holotype MGPT-PU 135300, tail valve, width 9.5 mm, Figs 143 A – D. Type locality. Montegibbio (Emilia-Romagna, Italy). Type stage. Miocene (Tortonian). Material examined. Miocene (Tortonian): Italy: Montegibbio: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF94F070FADF9786B6B90C5.taxon	description	Description. Head and intermediate valves unknown. Tail valve flat, characteristic bell shape at top, semicircular in lower part (WT / LT = 1.22), anterior margin slightly sinuous bordering a large and triangular jugal area, side margins sinuous, rounded posterior margin, mucro subcentral, not elevated and little evident, antemucronal and postmucronal slopes slightly convex. Tegmentum covered with large, dense and elevated granules of irregular shape, except on JA, becoming larger and coalescing towards margins, granules highly variable, from single granules, almost ellipsoidal to quadrangular, to coalescing granules fused to form irregular cords, especially near JA, where they are also less evident, top of granules convex; some granules close to posterior margin are rectangular in shape, 2 – 3 times larger than neighboring granules. Each granule with many megalaesthetes and micraesthetes (more than 10) arranged without any apparent order. Articulamentum thick, apophyses expanded with rounded edges, widely separated by an unconnected sinus, nine slits, deep and dorsally slightly elevated, forming a channel, teeth broad and roughed dorsally, very irregular.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF94F070FADF9786B6B90C5.taxon	discussion	Remarks. The fossil record of Craspedochiton brunettii Dell’Angelo, Giuntelli, Sosso & Zunino, 2016 is represented thus far by a single tail valve from the upper Miocene of Montegibbio (Dell’Angelo et al. 2016). This valve is very characteristic and different from the other European Craspedochiton spp. here discussed. The presence of the insertion lamina with numerous slits and granules with the top convex confirm the attribution to the genus Craspedochiton. Comparisons. The main difference with the tail valves of Craspedochiton altavillensis (Seguenza, 1876) regards the shape of the tegmentum, elliptical in C. altavillensis with the width about twice the length (WT / LT = 1.62 – 1.89), much more lengthened and with a characteristic bell shape at the top in C. brunettii (WT / LT = 1.22). Furthermore the jugal area is narrower in C. altavillensis, the valves are higher, and the granules are more regular.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF94F070FADF9786B6B90C5.taxon	distribution	Distribution. Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, Italy: Montegibbio (Dell’Angelo et al. 2016).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF84F080FADF9D86BBD9115.taxon	description	Fig. 144	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF84F080FADF9D86BBD9115.taxon	materials_examined	Type material. Holotype: MHNBx 2020.4.1, tail valve, width 2.8 mm, Figs 144 A – D. Type locality. Gamachot (France). Type stage. Miocene (Early Burdigalian). Material examined. Type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF84F080FADF9D86BBD9115.taxon	description	Description. Head and indeterminate valves unknown. Tail valve elliptical (WT / LT = 1.73), anterior margin straight, jugal area triangular, very large, mucro posterior, poorly developed, antemucronal slope slightly convex, postmucronal slope weakly concave. Tegmentum covered with small granules of irregular shape, coalescing to form irregular striae near side margins. Articulamentum thick, apophyses trapezoidal, widely separated by a large sinus, teeth broad, higly irregular, ten deep slits.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF84F080FADF9D86BBD9115.taxon	discussion	Remarks. Craspedochiton fontlevensis Dell’Angelo, Lesport, Cluzaud & Sosso, 2020, described upon a single tail valve, is only known from the lower Miocene of Noaillan (Dell’Angelo et al. 2020 b). The single valve available is not well preserved, but the characters of the shell are so distinctive to warrant a specific recognition. Fig. 144 B show the valve always in dorsal view but more inclined, in order to better highlight some characteristics of the mucro. Comparisons. See Tab. 23 for a comparison with the Craspedochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF84F080FADF9D86BBD9115.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Gamachot (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF74F090FADFBEB6B5991FF.taxon	description	Fig. 145	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF74F090FADFBEB6B5991FF.taxon	materials_examined	Type material. Holotype MHNBx 2020.2.8, tail valve, width 3 mm, Figs 145 F – H. Paratypes: MHNBx 2020.1.6, head valve, width 4 mm, Figs 145 A – B; MHNBx 2020.1.5, intermediate valve, width 4.5 mm, Figs 145 C – E. Type locality. Gaas, Lagouarde (France). Type stage. Oligocene (Rupelian). Material examined. Lower Oligocene (Rupelian): Gaas: type material plus 17 valves (AC, DA, JCV). Maximum width of the valves: 4 / 4.5 / 3.1 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF74F090FADFBEB6B5991FF.taxon	description	Description. Head valve semicircular, with five well evident radial ribs, slope almost straight. Intermediate valve trapezoidal (WT / LT = 1.56 – 1.78), semicarinate in anterior profile, elevated (H / W = 0.44 – 0.57), anterior margin almost straight in jugum, side margins rounded, posterior margin slightly concave at both sides of prominent apex, jugal area large, triangular, lateral areas not separated from central area. Tail valve broadly circular, mucro subcentral, not elevated and poorly developed, antemucronal and postmucronal slopes almost straight. Tegmentum covered with large, elevated granules of irregular shape, except on JA, furrowed by strong longitudinal cords (poorly delimited in JA abraded of tail valve), shape of granules highly variable, from single granules, regularly ellipsoidal, to coalescent granules, top of granules convex. Articulamentum with trapezoidal and widely separated apophyses, expanded in tail valves, insertion lamina reduced to narrow band in HV, large posterior callus in tail valves, slit formula 5 / 1 / 7, teeth broad, roughed dorsally, more irregular in tail valves, slits deep, dorsally slightly elevated, forming channel.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF74F090FADFBEB6B5991FF.taxon	discussion	Remarks. The fossil record of Craspedochiton lozoueti Dell’Angelo, Lesport, Cluzaud & Sosso, 2020 is limited to the lower Oligocene of Gaas (Dell’Angelo et al. 2020 b). This is the oldest species of Craspedochiton known from the Cenozoic of Europe. Comparisons. The sculpture of the jugal area of Craspedochiton lozoueti (furrowed by strong longitudinal cords is different from that of the other species of Craspedochiton studied here, granular (albeit with less evident granules in some cases) or smooth.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF74F090FADFBEB6B5991FF.taxon	distribution	Distribution. Lower Oligocene: northeastern Atlantic (Rupelian): Aquitaine Basin, France: Gaas (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF64F0A0FADFA866BBD9063.taxon	description	Fig. 146	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF64F0A0FADFA866BBD9063.taxon	materials_examined	Type material. Holotype MZB 50583, tail valve, width 3.8 mm, Figs 146 A – D. Type locality. Maureilhan (France). Type stage. Miocene (Lower Burdigalian). Material examined. Type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF64F0A0FADFA866BBD9063.taxon	description	Description. Head and intermediate valves unknown. Tail valve broadly circular (WT / LT = 1.10), anterior margin strongly convex, jugal area triangular, not raised, mucro subcentral, not elevated and poorly developed, antemucronal slope slightly convex, postmucronal slope straight. Tegmentum covered with small, roundish, elevated granules, irregularly disposed, only in jugal area giving impression of forming sinuous longitudinal striae. Articulamentum thick, expanded around posterior margin, apophyses trapezoidal, posterior margin not uniform, with a large posterior callus, and seven slits, two deep and strong and five less developed located in between them.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF64F0A0FADFA866BBD9063.taxon	discussion	Remarks. The species is known by a single tail valve, superficially similar to the genus Acanthochitona Gray, 1821, but with important differences concerning JA and the articulamentum. The jugal area, not raised and in continuity with the pleural areas, has the same type of sculpture formed by small, roundish and elevated granules, but arranged more regularly, giving the impression of forming sinuous longitudinal striae (Fig. 146 B); the profile of the valve is evenly rounded. The articulamentum is expanded around the posterior margin, which is not uniform throughout, with two protrusions and strong slits that, based on their position, correspond to the incisions in the genus Acanthochitona. A further five weaker incisions (much less developed and irregular) are placed between the primary slits, partially closed by a large back callus, that delimit four small, sharp teeth (Fig. 146 C). The number of slits can therefore be considered 7 (2 + 5). Šulc (1934: p. 16) described the new subgenus Pseudoacanthochiton for species characterized by tail valves similar to those of Acanthochitona, but with additional slits on the insertion plate. Considering that some tail valves of Acanthochitona with additional slits on the insertion plate had been reported [see above for A. fascicularis (Linnaeus, 1767)], we prefer to consider the genus Pseudoacanthochitona (Šulc, 1934) as a synonym of Acanthochitona Gray, 1821. Comparisons. See Tab. 22 and 23 for a comparison with the Acanthochitona and Craspedochiton spp. considered in this study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF64F0A0FADFA866BBD9063.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Maureilhan (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF54F0B0FADFA3D68E49361.taxon	description	Fig. 147	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF54F0B0FADFA3D68E49361.taxon	materials_examined	Type material. Holotype in Bałuk collection, reg. No. BkK-A 57, an intermediate valve (Bałuk 1971: pl. 6, fig. 9). Type locality. Korytnica, southern slopes of the Holy Cross Mts. (Poland). Type stage. Middle Miocene. Material examined. Miocene (Tortonian): Italy: Montegibbio: 28 valves (BD 1139, MZB 32121 – 32124, Figs 147 A – H), Rio di Bocca d’Asino: 3 valves (BD 1140). Maximum width of the valves: 1.5 / 1.9 / 1.8 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF54F0B0FADFA3D68E49361.taxon	description	Description. Head valve semicircular. Intermediate valve broadly rectangular (W / L = 1.50 – 1.83), carinate in anterior profile, anterior margin almost straight, side margins rounded, posterior margin almost straight to slightly concave at both sides of prominent apex, projecting slightly outside posterior margin, jugal area large, lateral areas slightly raised, poorly separated from central area. Tail valve semicircular, anterior margin straight, mucro subcentral, poorly developed. Tegmentum covered with densely spaced, small, flat, roundish / oval granules, arranged in arcuate rows, sometimes less visible on JA; granules fairly regular, with a central megalaesthete surrounded by 6 – 10 micraesthetes. Articulamentum only slightly larger than tegmentum in intermediate valves, trapezoidal and widely separated apophyses just more expanded in tail valves, insertion lamina reduced to narrow band in head valve, slit formula 5 / 1 / 6 – 10, teeth irregular, roughed dorsally.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF54F0B0FADFA3D68E49361.taxon	discussion	Remarks. This species was described by Bałuk (1971, 1984) upon many small valves from the Middle Miocene (Badenian) of Korytnica (Poland). The species was also reported by Laghi (1977), Dell’Angelo et al. (2016) from the Miocene (Tortonian) of N. Italy, and Ruman & Hudácková (2015) from the Middle Miocene of Slovakia. There is a certain variability in the shape of intermediate valves, with the apex sometimes very pronounced and projecting slightly outside posterior margin (e. g., Bałuk 1984, pl. 11, fig. 3). Comparisons. Craspedochiton minutulus Bałuk, 1971 differs from the other Craspedochiton spp. described here mainly by its more regular sculpture, with granules arranged in arcuate rows, different from the irregular granules of variable size, more or less coalescing, shown by the other Craspedochiton spp.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF54F0B0FADFA3D68E49361.taxon	distribution	Distribution. Middle Miocene: Paratethys (Langhian-Serravallian): Slovakia: Devínska Nová Ves, Rohožník (Ruman & Hudácková 2015), Poland: Korytnica, Lychów, Węglin, Weglinek (Bałuk 1971, 1984; Macioszczyk 1988). Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, Italy: Montegibbio, Rio di Bocca d’Asino (Laghi 1977; Dell’Angelo et al. 1999, 2016).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF44F0D0FADF93C690797F1.taxon	description	Fig. 148	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF44F0D0FADF93C690797F1.taxon	materials_examined	Type material. Syntype MPM 6090, head valve, coll. Doderlein, figured by Sacco (1897: pl. 7, fig. 38) and Laghi (1977: pl. 4, fig. 2), Fig. 148 A. Type locality. Montegibbio (Emilia-Romagna, Italy). Type stage. Miocene (Tortonian). Material examined. Lower Miocene: Italy: Sciolze: 1 head valve, BS. 105.04.001, width 10 mm, figured by Sacco (1897: pl. 7, fig. 33) and Laghi (1977: pl. 4, fig. 1). Maximum width of the head valves: 29 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF44F0D0FADF93C690797F1.taxon	description	Description. Intermediate and tail valves unknown. Head valve semicircular, large and strong, flat, with five well evident radial ribs which continue on the wide insertion lamina. Tegmentum covered with large granules of irregular oval shape. Articulamentum with insertion lamina particularly expanded, greater than length of tegmentum, with 5 deep incisions that divide insertion plate into 6 strong insertion teeth.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF44F0D0FADF93C690797F1.taxon	discussion	Remarks. Sacco (1897) described the new species Acantochiton (sic) costatus based on some head and tail valves from the Torino Hill (Sacco 1897: pl. 7, figs 33 - 37), although he considered doubtful conspecificity among those head plates. The head valves are characterized by the presence of large radial ribs, and especially by the insertion lamina particularly expanded. Only one of the three valves figured by Sacco is still present in the Bellardi-Sacco collection (Sacco: pl. 7, fig. 33, width 10 mm), whereas the other two valves from the Collection Rovasenda are missing. Sacco also described another head valve from Montegibbio as Acantochiton (sic) costatus var. mutinocrassa (Sacco: pl. 7, fig. 38); this specimen is identical to the head valves from the Torino Hill, but larger (width 22 mm). Craspedochiton costatus (Sacco, 1897) was considered a valid species by some authors (Laghi 1977; Bałuk 1984), or a synonym of other related species by others, e. g., C. profascicularis (Boettger, 1907) (Šulc 1934; Zilch 1934; Studencka & Dulai 2010) or C. altavillensis (Seguenza, 1876) (Dell’Angelo et al. 1999). Dell’Angelo et al. (2016) considered the remarkable expansion of the insertion lamina as an important character to differentiate these valves from C. altavillensis, where the insertion lamina is reduced to a rather narrow band that does not appear to vary with the valve’s size. The expansion of the insertion lamina in each of the two head valves from Montegibbio figured by Laghi (1977), the valve described by Sacco as A. costatus var. mutinocrassa (Laghi, pl. 4, fig. 2) and a still larger one (Laghi, pl. 4, fig. 3, width 29 mm) is greater than the length of the tegmentum (Figs 148 A – B). The taxon Acanthochiton costatus Adams & Angas, 1864 has priority over the fossil Acantochiton costatus Sacco, 1897, for which Dell’Angelo et al. (2016) proposed the name Craspedochiton mutinocrassus (Sacco, 1897). Therefore, only three head valves can be reliably attributed to Craspedochiton mutinocrassus: the valve from Sciolze (Sacco 1897, pl. 7, fig. 33; Laghi 1977, pl. 4, fig. 1; Ferrero Mortara et al. 1984, pl. 55, fig. 4; Figs 148 C – D) and the two valves from Montegibbio (Sacco 1897, pl. 7, fig. 38; Laghi 1977, pl. 4, figs 2 – 3; Figs 148 A – B). The tegmentum is generally abraded, only in one valve (MPM 6090, Fig. 148 A) is possible to see, in some area, an ornamentation with ellipsoidal granules (Laghi 1977, pl. 4, fig. 2 B). It is not possible to identify with certainty the tail valve figured by Sacco (1897, pl. 7, figs 36 – 37) as A. costatus, whose conspecificity with head valves was already considered doubtful by the same author. The shape of the tegmentum of the valves is different from that of C. altavillensis, and e ven the attribution to the genus Craspedochiton is uncertain. This valve is discussed and figured in the section dedicated to the “ Species of unclear taxonomic position ”, and it is left in open nomenclature as Craspedochiton? sp. Doderlein reported from Montegibbio one species of Polyplacophora [Chiton zibinicus (Doderlein, 1864) = Rhyssoplax olivacea (Spengler, 1797)], and also a species of barnacle (Coronula bifida Bronn, 1831), common within the Pliocene record of the Mediterranean (Dominici et al. 2011). Laghi (1977) made a provisional assignment of this finding to C. costatus, and we include here this taxon, that Laghi probably has seen during his work at Modena, studying the Doderlein collection. Comparisons. See Tab. 23 for a comparison with the Craspedochiton spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF44F0D0FADF93C690797F1.taxon	distribution	Distribution. Lower Miocene: Proto-Mediterranean Sea (Burdigalian): N. Italy: Sciolze (Sacco 1897; Laghi 1977; Dell’Angelo et al. 2016). Upper Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, N Italy: Montegibbio (Sacco 1897; Laghi 1977; Dell’Angelo et al. 2016).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF24F0E0FADFC8C69989698.taxon	description	Fig. 149	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF24F0E0FADFC8C69989698.taxon	materials_examined	Type material. Holotype NHMW 1863 / 00015 / 0861, tail valve, width 2.8 mm (Figs 149 A – D). Type locality. Niederleis (Austria). Type stage. Middle Miocene. Material examined. Middle Miocene: Austria: type material.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF24F0E0FADFC8C69989698.taxon	description	Description. Head valve unknown. Intermediate valves broadly rectangular, anterior margin almost straight, posterior margin with a prominent apex, jugal area large. Tail valve oval, anterior margin almost straight in jugum, mucro slightly posterior, poorly developed, antemucronal slope almost atraight, postmucronal slope weakly concave just under mucro. Tegmentum covered with elevated, roundish / oval irregularly arranged granules, coalescing towards anterior margin, except on JA, smooth; granules with many megalaesthetes and micraesthetes (up to 20 – 25 or more) arranged without any apparent order. Articulamentum with apophyses trapezoidal, not expanded, widely separated by a large sinus, teeth broad, roughed dorsally, slit formula - / 1 / 7.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF24F0E0FADFC8C69989698.taxon	discussion	Remarks. Craspedochiton schafferi (Šulc, 1934) was described upon a tail valve from Niederleis (Austria), and three intermediate valves from Knínice (Czech Republic. The tail valve (holotype) is deposited at the NHMW. The material from the Lower Badenian of Korytnica attributed to this species (Bałuk 1971) was later revised and attributed to Craspedochiton profascicularis (Boettger, 1907) by Bałuk (1984). Šulc (1934) reports a width of 4 mm, probably also considering both complete apophyses. Currently, part of the valve is missing (compare Kroh 2003: fig. 4 and Fig. 149 A), and the actual width is 2.8 mm. The three intermediate valves from Knínice are not present at NHMW; Šulc (1934) does not provide an exaustive description but provides only some indications These segments are relatively long, the articulamentum covers most of the shell and the ornamentation, similar to the tail valve, consists of irregular protuberances. On each side of an incision occurs), and an illustration (Šulc 1934: pl. 1, fig. 22), insufficient to define features. Another tail valve was recently described by Dulai (2024 b) from Letkés (Hungary), making it the first known record of C. schafferi since the original description by Šulc (1934). Šulc attributed this species to the genus Cryptoconchus Burrow, 1815, a genus characterized by the tegmentum reduced to a small jugal area on each valve (Van Belle 1983: 140), and not consistent with the characters of our valve. The attribution to the genus Notoplax H. Adams, 1862, is not consistent, following Gowlett-Holmes (1991) who discussed the status of the relations between the genera Notoplax and Craspedochiton Shuttleworth, 1853, and redefined the genus Notoplax, and so we attribute the Šulc’s species to the genus Craspedochiton. Comparisons. Craspedochiton schafferi shows some similarity with C. brunettii Dell’Angelo, Giuntelli, Sosso & Zunino, 2016, but it differs from the shape of the tegmentum of the tail valve (without the characteristic bell shape at the top), and the more regular shape of the granules, ellipsoidal, size fairly constant and less dense.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF24F0E0FADFC8C69989698.taxon	distribution	Distribution. Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria: Niederleis; Czech Republic: Knínice (Šulc 1934; Kroh 2003); Hungary: Letkés (Dulai 2025 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF14F0E0FADFC646B079310.taxon	type_taxon	Type species. Chiton larvaeformis de Blainville in Burrow, 1815, by subsequent designation (Gray 1821).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF14F0E0FADFC646B079310.taxon	distribution	Distribution. Cryptoplax is known from the Miocene to the Recent, with a living distribution in tropical and temperate Indian and central western Pacific Oceans and the Red Sea (Gowlett-Holmes 2001). The fossil record extends back to the Miocene in Europe (Laghi 1977; Dell’Angelo et al. 1999, 2016; Dulai 2001, 2005; Ruman & Hudáčková 2015), Melanesia (Schwabe et al. 2008) and W. Pacific Islands (Ladd 1966), the Miocene-Pliocene in Australia (Cotton 1964) and the Pleistocene in the Red Sea (Selli 1944, 1973; Dell’Angelo et al. 2020 a).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF14F0E0FADFC646B079310.taxon	discussion	Remarks. Three species of Cryptoplax are known from the Miocene of N. Italy and Paratethys: C. weinlandi Šulc, 1934, C. lanceolatus Laghi, 1977, and C. margitae Dulai, 2001. Chitonellus gigas O. G. Costa (1854), described upon an intermediate valve from the Pleistocene of Cannitello (Reggio Calabria), does not, in fact, belong to the genus Cryptoplax (= Chitonellus Lamarck, 1819); the generic attribution of Costa’s species remains uncertain (see below). The main characters of the species here discussed are reported in Tab. 24.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF04F300FADFAE86BC690A9.taxon	description	Fig. 150	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF04F300FADFAE86BC690A9.taxon	materials_examined	Type material. MPUM 18972 (one head and one tail valve), Figs 150 A – B. Type locality. Montegibbio (Piedmont, Italy). Type stage. Miocene (Tortonian). Material examined. Upper Miocene: Italy (Tortonian): Montegibbio: 1 intermediate valve (MZB 32131, Figs 150 C – D). Maximum width of the valves: 1.5 / 1.6 / 1.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF04F300FADFAE86BC690A9.taxon	description	Description. Valves longer than wide. Head valve trapezoidal. Intermediate valve pentagonal, jugal area rectangular, converging in apex. Tail valve polygonal, jugal area smooth, on which are seen, aligned in parallel rows, exit holes of aesthete channels. HV sculptured with three trapezoidal arcuate ribs, centered in apex, of increasing size, and remains of weak lumpy ribs, aligned forward, only visible on left side of HV; PLA ornamented with weak lumpy ribs aligned forward. Articulamentum with apophyses elongated triangular in outline in intermediate valves, head valve with apical area shaping a circular segment, slits not visible on insertion plate of head valve, slit formula? / 0 / 0.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF04F300FADFAE86BC690A9.taxon	discussion	Remarks. Cryptoplax lanceolatus Laghi, 1977 was described on the basis of a head and tail valve from Montegibbio, not too well preserved, with a sculpture of weak lumpy ribs (the “ deboli coste grumose ” in Laghi 1977) quite different in form from the longitudinal, slightly undulated ribs typical of C. weinlandi Šulc, 1934. Later, a single intermediate valve, also from Montegibbio, has been attributed to this species by Dell’Angelo et al. (2016). The head valve depicted by Laghi is almost completely decorticated, the narrow insertion plate does not appear engraved, perhaps because the teeth are completely worn. Comparisons. C. margitae Dulai, 2001, is the species whose sculpture is closer to Cryptoplax lanceolatus (see below).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEF04F300FADFAE86BC690A9.taxon	distribution	Distribution. Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin, Italy: Montegibbio (Laghi 1977; Dell’Angelo et al. 2016).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECF4F320FADFA746F489108.taxon	description	Fig. 151	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECF4F320FADFA746F489108.taxon	materials_examined	Type material. Holotype: HNHM M. 99.113, intermediate valve, width 1.16 mm. Type locality. Borehole Szokolya- 2, 92.7 - 93 m, Börzsöny Mts (Hungary). Type stage. Middle Miocene, marly sandstone. Material examined. Lower Miocene (Burdigalian): France: Carrière Vives: 1 head valve (MZB 50586, Fig. 151 A); Gamachot: 3 intermediate valves (BD 1141, MZB 50587, Fig. 151 G). Middle Miocene: Romania: Kostej: 1 valve (NHMW 1869 / 0001 / 0797, as Cryptoplax weinlandi, Figs 151 B – C); Lăpugiu de Sus: 1 valve (NHMW 2010 / 0256 / 0031), Figs 151 D – F; Hungary: Bánd: 2 intermediate valves (BD 1142, Fig. 151 H). Maximum width of the valves: 1.3 / 1.7 / -.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECF4F320FADFA746F489108.taxon	description	Description. Tail valve unknown. Valves longer than wide. Head valve horseshoe shaped, very small, anterior margin rounded, posteriorly straight, with oval area near apex, smooth and only covered by an irregular arrangement of megalaesthete and micrasthetes. Intermediate valve of pentagonal shape, very small, posterior margin obtusely angled. Jugal area smooth, rectangular, with edges slightly undulated. Tegmentum strongly porous, HV sculptured with irregularly disposed granules, roundish near the posterior margin, becoming larger, more elongate, and tending to coalesce towards anterior margin; PLA ornamented by 4 – 5 longitudinal rows of rounded granules, parallel to each other and sides of JA, granules roundish, elevated, well separated near apex, and tending to coalesce towards anterior valve margin. Each granule with a central megalaesthete and several micraesthetes (6 - 8 or more irregularly spaced all along in HV; many in PLA irregularly arranged, mainly on same side, some micraesthete can be seen also in grooves between rows of granules; an irregular arrangement of micraesthetes is present on JA too). Articulamentum with apophyses relatively short, ca ¼ of whole length of valve, of triangular shape, slit formula 3 / 0 / 0.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECF4F320FADFA746F489108.taxon	discussion	Remarks. Cryptoplax margitae Dulai, 2001 was described from two intermediate valves of the Middle Miocene (Badenian) of Hungary (Dulai 2001, 2005). Other 4 valves (one head, partly eroded, and 3 intermediate) from the lower Miocene (Burdigalian) of the Aquitaine Basin were attributed to this species by Dell’Angelo et al. (2020 b); two valves (1 head, well preserved, and 1 intermediate) were identified in the Šulc collection deposited at NHMW; two intermediate valves from the Middle Miocene of Bánd (Hungary) are added here. These latest findings allow to include the description of the head valve, while the tail one is still unknown. The valves of Cryptoplax margitae are also very similar to Choneplax indica Odhner, 1919, an extant species of the family Acanthochitonidae, known from the western Indian Ocean: Mozambique Channell, Madagascar, Réunion, Mauritius, Rodrigues, and Arabian Sea (Dell’Angelo et al. 2004; Dinapoli & Janssen 2009). The analysis of figures of this species (Kaas 1986: figs 67 – 72; Dell’Angelo et al. 2004: pl. 6, figs 8 – 13; Schwabe 2004: fig. 11 A – D; Dinapoli & Janssen 2009: pl. 16, figs b – c) confirms indeed their resemblance. The small dimensions, shape of head and intermediate valves, the smooth JA, the sculpture and shape of granules irregularly disposed in HV and arranged in 4 – 5 longitudinal rows in PLA are really very similar, if not identical. This similarity was already noticed by Dulai (2005: p. 42), while discussing intermediate valves II of Cryptoplax weinlandi, which does not present the typical granulated sculpture characteristic of C. margitae. The genus Choneplax Carpenter in Dall, 1882, includes four Recent species (Sirenko 2003): C. lata (Guilding, 1829) from the Caribbean, C. hastata (Sowerby, 1840), habitat unknown (fide Sirenko 2003), C. indica Odhner, 1919 from the W. Indian Ocean, and C. littlerorum Sirenko, 2003, from American Samoa. Choneplax is known from the Pleistocene of the Red Sea (Dell’Angelo et al. 2020 a). Before the question of a reassignment of Cryptoplax margitae to Choneplax is taken into consideration, the relation between extant Cryptoplax and Choneplax needs to be clarified. Consequently, we prefer to adhere to Dulai’s attribution of margitae to Cryptoplax. Choneplax (attributed to the family Acanthochitonidae, see Sirenko 2006 and Irisarri et al. 2014) differs from Cryptoplax in the strong imbrications of all the valves, and most importantly in the different characters of girdle. Entire live collected individuals of both species are easily separable, while it is more difficult to separate loose valves. Sirenko (2003) suggests that the evolution of the superfamily Cryptoplacoidea (which includes three families: Acanthochitonidae Pilsbry, 1893, Cryptoplacidae H. & A. Adams, 1858, and Hemiarthridae Sirenko, 1997) can be connected with a reduction of the tegmentum and, as a result, a reduction of slits of the insertion plates. Comparisons. Cryptoplax lanceolatus Laghi, 1977 is the species whose sculpture most closely resembles that of Cryptoplax margitae (see above). However, the sculpture in C. lanceolatus is more rugged, consisting of weak, lumpy ribs, in contrast to the rows of roundish, elevated, and more regular granules seen in C. margitae. Moreover, the head valve differs — C. lanceolatus displays three arcuate ribs — and so does the outline of the intermediate valves (compare Figs. 150 C with 151 D and 151 G).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECF4F320FADFA746F489108.taxon	distribution	Distribution. Lower Miocene: northeastern Atlantic (Burdigalian): Aquitaine Basin, France: Carrière Vives, Gamachot (Dell’Angelo et al. 2020 b); Middle Miocene: Central Paratethys (Langhian-Serravallian): Hungary: Bánd, Szokolya- 2 borehole (Dulai, 2001, 2005; this study), Romania: Kostej, Lăpugiu de Sus (this study).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECD4F340FADFB146FED92C6.taxon	description	Fig. 152	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECD4F340FADFB146FED92C6.taxon	materials_examined	Type material. Unknown, not present in NHMW. Type locality. Rudoltice (Czech Republic). Type stage. Middle Miocene. Material examined. Middle Miocene: Central Paratethys (Langhian-Serravallian): Austria: Enzesfeld: 5 valves (BD 1143), Forchtenau: 40 valves (NHMW 1866 / 0001 / 1210, Figs 152 D – F), Niederleis: 22 valves (NHMW 1863 / 0058 / 0033, 1866 / 0001 / 0977), Steinabrunn: 1 valve (NHMW 1871 / 0010 / 0410), Romania: Kostej: 5 valves (BD 1144, NHMW 1869 / 0001 / 0797), Lăpugiu de Sus: 233 valves (BD 1145, Figs 152 G – H, NHMW 1868 / 0001 / 0634, 1876 A / 0011 / 0174, 2010 / 0256 / 0012); Hungary: Bánd: 161 valves (BD 1146), Letkés: 63 valves (BD 1147), Márkháza: 2 valves (BD 1148); Eastern Paratethys: Ukraine: Horodok: 1 valve (BD 1149); Italy (Serravallian): Monchio di Sarzano Casina: 2 valves (BD 1150). Upper Miocene (Tortonian): Italy: Borelli: 2 valves (MZB 32130, Fig. 152 I, PG), Montegibbio: 14 valves (BD 1151), Rio di Bocca d’Asino: 83 valves (BD 1152, PG), Vargo: 1 valve (PG), Vigoleno: 29 valves (BD 1153), Villa Monti: 426 valves (BD 1154, Fig. 152 J, MZB 32125 – 32129, Figs 152 A – C, 152 K – M, PG). Maximum width of the valves: 6 / 9.8 / 8.5 mm.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECD4F340FADFB146FED92C6.taxon	description	Description. Head valve elongated semicircular, slope convex, Intermediate valves ii of pentagonal shape, smaller, length varying between 2.3 and 4.3 mm (WT / LT = 0.64 – 0.78), apex triangular, jugal area smooth, trapezoidal, with wider anterior base and wedge shaped sides; intermediate valves iii-vii larger and more elongated, length varying between 3.6 and 6.8 mm (WT / LT = 0.35 – 0.55), apex rounded or more pointed, jugal area smooth, rectangular, with parallel sides or slightly wedge-shaped. Tail valve of pentagonal shape, with low mucro at posterior end, antemucronal slope convex, postmucronal slope straight. Tegmentum strongly porous, with megalaesthetes and micraesthetes irregularly arranged in grooves between ribs; if ornamentation is granulose, most aesthetes are on granules, with a megalaesthete subcentral and micraesthetes irregularly around; HV ornamented with irregular wavy ribs emanating from posterior margin, joining in irregularly lobed network, dominated by radial direction, ribs may become granulose near apex; PLA of intermediate valves ii ornamented by up to 10 slightly undulating ribs, ribs may become granulöse near apex; PLA of intermediate valves iii-vii ornamented by 6 – 8 longitudinal, sometimes slightly undulating ribs, ribs may become granulöse near apex; tail valve as for intermediate valves iii-vii. Articulamentum with apophyses elongated triangular in outline, with very variable length, changing between 39 – 70 % of the total length; posterior end of insertion plate only slightly expanded posteriorly in tail valve; slits shallow, while middle is slightly deeper, slit formula 3 / 0 / 0.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECD4F340FADFB146FED92C6.taxon	discussion	Remarks. Cryptoplax weinlandi Šulc, 1934 was first mentioned by Boettger (1902) as to be described by Rolle; as reported by Šulc (1934) and Zilch (1934), the description of this species was never published by Rolle (the name was known only from his unpublished manuscript), and, therefore, the authorship pertains to Šulc (1934). As pointed out by Šulc (1934), the shape and appearance of the intermediate valves of C. weinlandi is variable. Some valves are characterized by an almost equal length and width and by more longitudinal, slightly undulated ribs in PLA, up to 10 per side (Figs 152 D – F, 152 I) and are interpreted as the intermediate valve ii, others are large sized, elongate triangular in outline, with less ribs in PLA, and are interpreted as the intermediate valve iii-vii (Figs 152 G – H). A detailed discussion of the length-width relationships of intermediate valves in C. weinlandi from Hungary and Romania is found in Dulai (2005) and Dell’Angelo et al. (2007 a). The intermediate valves show a great variability in the tegmentum sculpture: more regular longitudinal ribs (Fig. 152 G) or irregular and wavy (Fig. 152 D), and also the size of the articulamentum in tail valves is variable (Dell’Angelo et al. 2016, pl. 7, figs 23, 25). A new species name (Cryptoplax viciani), based on intermediate valves from Lăpugiu de Sus differing from C. weinlandi in morphological characters, was presented by Dell’Angelo et al. (2005) as a congress abstract, and must be considered nomen nudum. Dell’Angelo et al. (2007 a) proved that these shorter intermediate valves represented the valve ii of C. weinlandi. Comparisons. The ornamentation of the valves of Cryptoplax weinlandi (longitudinal ribs) is basically different from the more or less granulose ornamentation of the two other species of Cryptoplax here considered, C. lanceolatus Laghi, 1977 and C. margitae Dulai, 2001. See Tab. 24 for a comparison with the Cryptoplax spp. considered in the present study.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECD4F340FADFB146FED92C6.taxon	distribution	Distribution. Middle Miocene: Proto-Mediterranean Sea (Serravallian): N. Italy: Monchio di Sarzano Casina (Dell’Angelo et al. 2016). Central Paratethys (Langhian-Serravallian): Austria: Enzesfeld, Forchtenau, Niederleis, Nodendorf, Steinabrunn (Šulc 1934; Zilch 1934; Sieber 1956; Kroh 2003; this study); Czech Republic: Borač, Knínice, Lysice, Porzteich, Rudoltice, Sudice, Židlochovice, Vranovice (Šulc 1934; Zilch 1934; Zágoršek 2006); Slovakia: Devínska Nová Ves, Kúty (Ruman & Hudácková 2015); Poland: Korytnica (Baluk 1971, 1984); Hungary: Bánd, Devecser, Letkés, Márkháza, Szokolya, Várpalota (Dulai 2001, 2005, 2025 a, 2025 b; Dulai & Katona 2024; this study); Romania: Băseşti, Bujturi, Coştei, Lăpugiu (Šulc 1934; Zilch 1934; Marinescu 1964; Dell’Angelo et al. 2007 a; this study); Eastern Paratehys: Ukraine: Horodok (this study). Upper Miocene: Proto- Mediterranean Sea (Tortonian): Po Basin, Italy: Borelli, Montegibbio, Rio di Bocca d’Asino, Vargo, Vigoleno, Villa Monti (Laghi 1977; Dell’Angelo et al. 1999, 2016); S. Italy (Messinian): Petralia Sottana (Sicily) (Ruggieri 1982 a). Species of unclear taxonomic position All the species of unclear taxonomic position present in the literature, or whose presence in the fossil state is not convincigly confirmed, are briefly discussed here. These 19 species are not reported in a systematic order, but listed by the year of publication (from 1797 to 1897), with some original figures included in a summary plate (Fig. 153).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FECA4F360FADF9336B3C9458.taxon	discussion	Remarks. Morais (1941) reported Chaetopleura fulva (Wood, 1815), now accepted as Chaetopleura angulata Spengler, 1797, in a list of molluscs from the Pliocene of Marinha Grande (Guarda Nova and Matos), Portugal. The fossils were identified by Reginald Cox, palaeontologist at the British Museum (Natural History), but unfortunately, no description or illustration of the valve (s) were given. According to Morais (1941), the malacofauna from Marinha Grande was presented to the Minerological and Geological Museum of the University of Coimbra, but it was impossible to locate such material (Dell’Angelo & Silva 2003: p. 9). Chaetopleura angulata lives along the West coast of the Iberian Peninsula, as as well as in South America from Cabo Frio, Brasil, to Cape Horn (Kaas & Van Belle 1988). The fossil record is known from the Pleistocene of Uruguay (Figueiras 1961) and a single record from the Pliocene of Portugal (Morais 1941), not further confirmed by later studies.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC94F370FADFE24687A9622.taxon	discussion	Remarks. Kaas (1985 a) provided an accurate analysis of the taxonomic history of the three European species of Acanthochitona currently accepted at the time, i. e. A. fascicularis (Linnaeus, 1767), A. crinita (Pennant, 1777) and A. discrepans (Brown, 1827), discussing also their various interpretations in the past. Before, A. fascicularis was reported mainly as A. communis Risso, 1826, or A. discrepans (non Brown), and so all the references reported as A. discrepans from Italian localities (and from few other Mediterranean localities) are attributable to A. fascicularis in our conception (see above). Some reports of A. discrepans from the English Pliocene (Coralline Crag, Sutton) could be related to the true A. discrepans, a species still living at present along the British coast (Kaas 1985 a; Vončina et al. 2023). Acanthochitona discrepans is very similar to A. crinita, with the main differences concerning the number, size and arrangement of aesthetes on tegmental granules, and especially the girdle formations (Vončina et al. 2023), not appreciable in single valves only. Even the description and the figure provided by Wood (1872 / 1874) do not permit a certain specific attribution; so, we report A. discrepans in this appendix which groups together species whose fossil findings are not correctly attributable.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC94F370FADFE24687A9622.taxon	discussion	Remarks. Woodward (1833) reports Chiton octovalvis referring to Parkinson (1811: 44 “ Org. Rem. iii, t. 5, f. 5 ” Ibid. (Thorpe) One valve only ”), but Parkinson only note the finding of several separated valves at Grignon, near Paris (by Lamarck 1802) and figure a complete species from Grignon (reproduced here, Fig. 153 A), confirming the opinion of Lamarck, as to the number of its valves, which are eight [Lamarck (1802) described Chiton grinionensis from the Eocene of Grignon (Paris Basin) as: “ Chiton (Grinionensis) octovalvis? ”]. Later authors still considered Chiton octovalvis as a valid taxon, until Wood (1872 – 1874) observation that the species (“ a single valve from Thorpe ”) was not present in any known collection. This taxon should be, therefore, considered a species inquirenda. “ Chiton ” angulosus S. Wood, 1842	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC94F370FADFE24687A9622.taxon	discussion	Remarks. This species is reported by Wood (1842) from the Pliocene English Crag, without description. It has to be considered as a nomen nudum. “ Chiton ” tenuisculptus S. Wood, 1842	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC94F370FADFE24687A9622.taxon	discussion	Remarks. This species is reported by Wood (1842) from the Pliocene English Crag, without description. It has to be considered as a nomen nudum. “ Chiton ” subapenninus (Cantraine MS) de Ryckholt, 1845	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC94F370FADFE24687A9622.taxon	discussion	Remarks. This species is reported for the first time by de Ryckholt (1845), from the “ Upper Tertiary ” (Italy), and afterwards by Baily (1859; 1860 a; 1860 b; 1860 c) that considered it conspecific with Chiton miocenicus (Michelotti). This species is only mentioned in the literature without description. It is not reported by Cantraine (1835), and has to be considered a nomen nudum.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC84F370FADFCE26E9692FF.taxon	discussion	Remarks. Chitonellus gigas O. G. Costa, 1856, described on the base of an intermediate valve from the Pleistocene of Cannitello (Reggio Calabria, Italy) cannot be referred to the genus Cryptoplax (= Chitonellus Lamarck, 1819), and its generic attribution remains uncertain. The species is not present in the Costa’s collection hosted in the Paleontological Museum of Napoli University. The taxon “ Chitonellus ” gigas must be considered a nomen dubium (Dell’Angelo et al. 2007 a: p. 47). “ Lepidopleurus ” daubrei de Rochebrune, 1882 Figs 153 B – C	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC84F370FADFCE26E9692FF.taxon	discussion	Remarks. This species has been studied by Dell’Angelo et al. (2020 b), based upon a lot of six valves (5 intermediate and 1 tail) in the Benoist collection at MHNBx (MHNBx 2014.14.32.1 to 2014.14.32.3, MHNBx 2014.14.23.4 to 2014.14.23.6), labeled Lepidopleurus daubrei, from the Oligocene (Rupelian) of Gaas, France. De Rochebrune (1882) reported in his original description the locality of Lepidopleurus daubrei as Mérignac, near Bordeaux. However, Benoist (1882 a: p. xxix) reported that the three species sent to de Rochebrune (Tonicia gaasensis de Rochebrune, 1882, T. waltebledi de Rochebrune, 1882 and Lepidopleurus daubrei de Rochebrune, 1882), were from Gaas, as in the label in the Benoist collection (Dell’Angelo et al. 2020 b, figs 4 A – B). The description and figure given by de Rochebrune (Figs 153 B – C) do not match with the valves deposited at MHNBx; as discussed by Dell’Angelo et al. (2020 b), the six valves in the Benoist collection have been excluded from the type series and attributed to other Lepidochitona spp. (Dell’Angelo et al. 2020 b: tab. 2). Therefore, the taxon Lepidopleurus daubrei, whose generic attribution remains uncertain, has been considered as a nomen dubium (Dell’Angelo et al. 2020 b).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F380FADFF056B2396AD.taxon	description	Fig. 153 D	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F380FADFF056B2396AD.taxon	discussion	Remarks. This species has been studied by Dell’Angelo et al. (2020 b), that found a lot with two intermediate valves in the Benoist collection at MHNBx (MHNBx 2014.14.21.1 to 2014.14.21.2), labeled Tonicia wattebledi, from the Oligocene (Rupelian) of Gaas, France. This species was reported both as waltebledi or wattebledi, but the correct taxon is wattebledi (Dell’Angelo et al. 2020 b). The description and figure given by de Rochebrune (Fig. 153 D) do not match with the syntypes deposited at MHNBx (Dell’Angelo et al., 2020 b), the two intermediate valves in the Benoist collection have been excluded from the type series and attributed to other Lepidochitona spp. (L. larratensis Dell’Angelo, Lesport, Cluzaud & Sosso, 2020, see Dell’Angelo et al. 2020 b: tab. 2). The generic attribution of this species remains unassessed, and Tonicia wattebledi has been considered nomen dubium (Dell’Angelo et al. 2020 b). Unfortunately, de Rochebrune reports an incorrect reference for the figure of Tonicia waltebledi, “ pl. 3 fig. 2 ”, which corresponds instead to Tonicia pustulifera de Rochebrune, 1882, as reported in the caption of the table, and Dell’Angelo et al. (2020 b) include also the wrong figure in their paper (the correct figure is that of pl. 3 fig. 3 of de Rochebrune). We maintain, however, the opinion by Dell’Angelo et al. (2020 b) who consider Tonicia wattebledi as a nomen dubium.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F390FADFC706E279008.taxon	description	Fig. 153 E	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F390FADFC706E279008.taxon	discussion	Remarks. This species has been studied by Dell’Angelo et al. (2020 b), that found a lot with four intermediate valves in the Benoist collection at MHNBx (MHNBx 2014.14.23.1 to 2014.14.23.3, and MHNBx 2014.14.22), labeled Tonicia gaasensis, from the Oligocene (Rupelian) of Gaas, France. The description and figure given by de Rochebrune (Fig. 153 E) do not match with the syntypes at MHNBx (Dell’Angelo et al. (2020 b), the four intermediate valves in the Benoist collection have been excluded from the type series and attributed to other Lepidochitona spp. (Dell’Angelo et al. 2020 b: tab. 2). The generic attribution of this species remains unassessed, and the taxon Tonicia gaasensis has been considered as nomen dubium because the type material was not found (Dell’Angelo et al. 2020 b). “ Tonicia ” pustulifera de Rochebrune, 1882 Fig. 153 F	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F390FADFC706E279008.taxon	discussion	Remarks. Some of the species described by de Rochebrune (1882), including the species discussed here and the following three, are reported from the locality of “ Goubesville ”, referred to the Miocene. We do not know this locality, which could be “ Gourbesville ”, a well know site from the “ Redonian ” of the northwestern France, now considered upper Piacenzian – lower Gelasian in age (Lauriat-Rage 1981; Van Dingenen et al. 2015). The description and figure given by de Rochebrune (Fig. 153 F) preclude the identification of this species in the absence of the examination of the type material. We consider this taxon species inquirenda, whose generic status is uncertain.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F390FADFC706E279008.taxon	discussion	Remarks. De Rochebrune described intermediate and tail valves from Goubesville (now Gourbesville, see above). The description and figure given by de Rochebrune (Figs 153 G – H) are insufficient to identify the species, in the absence of the examination of the type material. This taxon is considered species inquirenda, and the generic attribution uncertain. “ Gymnoplax ” chalmasi de Rochebrune, 1882 Figs 153 I – J	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F390FADFC706E279008.taxon	discussion	Remarks. De Rochebrune described intermediate and tail valves from Goubesville (now Gourbesville, see above); there are no further citations for this taxon since the original description. The description and figure given by de Rochebrune (Figs 153 I – J) are insufficient to characterize the species, in the absence of the examination of the type material, we therefore consider this taxon as species inquirenda, whose generic attribution remains uncertain. “ Acanthochites ” meunieri de Rochebrune, 1882 Fig. 153 K	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC74F390FADFC706E279008.taxon	discussion	Remarks. De Rochebrune described intermediate and tail valves from Goubesville (now Gourbesville, see above). The description and figure given by de Rochebrune (Fig. 153 K) are insufficient to characterize the species, in the absence of the examination of the type material, and there have been no further reports after the original description. Weconsider this taxon as species inquirenda, whose generic status remains uncertain.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC64F390FADFA15694C92CD.taxon	description	Fig. 153 L	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC64F390FADFA15694C92CD.taxon	discussion	Remarks. De Rochebrune (1882) described a species of Acanthochitona from the Miocene of France (Serravallian), A. dulignoni, known for a single tail valve from Largileyre (Gironde, France) reported by Benoist (1882). Cossmann & Peyrot (1919) consider this valve close to Acanthochiton costatus Sacco, 1897 (now Craspedochiton mutinocrassus [Sacco, 1897], see Dell’Angelo et al. 2016: 90), refering to the valve figured by Sacco (1897: pl. 7, fig. 36), considered as Craspedochiton? sp. by Dell’Angelo et al. (2016: 91, pl. 6, figs 16 – 19). Dell’Angelo et al. (1999) considered this valve conspecific with Acanthochitona faluniensis (de Rochebrune, 1882) [considered = A. fascicularis (Linnaeus, 1767)]. Unfortunately, the description and figure of de Rochebrune (1882) (Fig. 153 L) and Cossmann & Peyrot (1919) are not sufficient to identify the species with certainty, and the type material of A. dulignoni is not present in the Benoist collection deposited at MHNBx (Charles Laurent, pers. com.). Therefore, A. dulignoni is considered nomen dubium.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC54F3A0FADFF056F4991EF.taxon	description	Fig. 153 M	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC54F3A0FADFF056F4991EF.taxon	discussion	Remarks. Wood (1848) described as Chiton fascicularis (?) some valves from the Pliocene Crag of Sutton (U. K.), with a sculpture similar to that of Acanthochitona fascicularis (Linnaeus, 1767) [now A. crinita (Pennant, 1777), after Kaas 1985 a], but more elevated and angular. De Rochebrune (1882) considered this material very different from the known species and proposed the new taxon Acanthochites woodi. The description and figure of Wood (1848) (reproduced here, Fig. 153 M) are not sufficient to identify the species with certainty, and the location of the Wood’s material is unknown, so this taxon should therefore be considered a species inquirenda. “ Chiton ” damesi Koenen, 1892 Fig. 153 N	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC54F3A0FADFF056F4991EF.taxon	materials_examined	Type material. Holotype: Gottinger Geologischen Institut (Koenen coll.), the tail valve figured by Koenen (1892: figs 23 a – c). Type locality. Latdorf, Magdeburg, eastern Germany (Lower Oligocene).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC54F3A0FADFF056F4991EF.taxon	discussion	Remarks. No further material of this Oligocene taxon has been found since its original description. A possible attribution to Leptochiton by Janssen (1978). was based on Koenen’s diagnosis, as the holotype was never restudied after. The main features deduced from the original description and figure (Fig. 153 N), based on a single tail valve, are: depressed valve; sculpture given by thin ribs arranged in groups with discontinuous orientation; absence of tubercles; presence on the ventral side of the valve of a longitudinal furrow which extends from the mucro to the anterior edge; apophyses connected by a lamina. These characters are sufficient to exclude the attribution of this species to the family Leptochitonidae. At present, the generic attribution of this species remains uncertain.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC54F3B0FADFAB66BCF97A1.taxon	discussion	Remarks. Rzehak (1893) described the new species Chiton reussi from the Lower Badenian of Oslawan (Czech Republic), based on a single incomplete intermediate valve, without providing an illustration. Instead Rzehak refers to the illustration of C. siculus Gray in Reuss (1860: pl. 8, figs 1 – 3), although pointing out that the ornamentations is slightly different). Therefore, it is impossible to confirm the attribution of this intermediate valve in the absence of the original specimen, and this taxon should be considered a species inquirenda. “ Acantochiton ” (sic) costatus Sacco, 1897 Fig. 153 O – P	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC54F3B0FADFAB66BCF97A1.taxon	discussion	Remarks. Sacco (1897) described the new species Acantochiton (sic) costatus based on some head and tail valves from the Turin Hill Miocene. The head valves (Sacco: pl. 7, figs 33 – 35) are now attributed to Craspedochiton mutinocrassus (Sacco, 1897) by Dell’Angelo et al. 2016 (see above), while the cospecificity of the tail valve (Sacco 1897: pl. 7, figs 36 – 37) was already considered doubtful by the same author. As evidenced by Dell’Angelo et al. (2016), the shape of the tail valve is different from that of C. altavillensis (more elliptical), as also the form of the apophysis. Even the attribution to the genus Craspedochiton is uncertain, since the articulamentum does not present the large teeth typical of this genus. Our inspection of the valve from the Bellardi-Sacco collection (MRSN, BS. 105.04.002, reproduced here, Figs 153 O – P) revealed only a couple of incisions, which may also correspond to those typical of the genus Acanthochitona, but we cannot exclude that other incisions have been obliterated due to erosion or missing due to the incompleteness of the posterior margin of the valve. We prefer to keep the previously described valve in open nomenclature as Craspedochiton? sp. Species of problematic assignment We list here, in systematic order and year of publication, those records whose determination is problematic, generally because of the scarcity of available material or its poor preservation state. Some original figures are included in the summary plate (Fig. 154).	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC14F3E0FADFEFC6B5B97D1.taxon	description	Fig. 154 H	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC14F3E0FADFEFC6B5B97D1.taxon	discussion	Remarks. A few small valves from the lower Pliocene of Liguria (Rio S. Antonino, Caranchi and Sestri Ponente), completely smooth and glossy, plus a tail valve from Caranchi (Fig. 154 H). At a first glance, the species appears similar to Rhyssoplax capecchii (Chirli, 2004); however, it differs in having a smooth tegmentum, whilst very fine radial and longitudinal striae always occur in C. capecchii. The teeth in the insertion plates are worn, so it is not possible to assess whether they are pectinated or not. We suggest a provisional generic assignment to Ischnochiton.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC04F3F0FADFDD36F439683.taxon	description	Fig. 154 Q	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
03FEF726FEC04F3F0FADFDD36F439683.taxon	discussion	Remarks. Four intermediate valves from the Pliocene of “ La Tagliata ” (Italy) (MPM n ° 1733, max width 3 mm, one of which reproduced here, Fig. 154 Q), that differ from the Acanthochitona spp. here examined mainly for many aspects: the outline of the valve (“ anterior margin semicircular ”), similar to the outline of the valve figured in Dell’Angelo & Smriglio (1999: fig. 9 A), but even more “ rounded ”; the jugal area not striated longitudinally (as the species of Acanthochitona from the upper Miocene of Saint-Clément-de-la-Place discussed above); possibly by the different shape of the granules, although the low quality Laghi’s figures prevent their accurate evaluation.	en	Dell’Angelo, Bruno, Sosso, Maurizio, Taviani, Marco (2025): The Cenozoic European Polyplacophora (Mollusca). Zootaxa 5704 (1): 1-377, DOI: 10.11646/zootaxa.5704.1.1, URL: https://doi.org/10.11646/zootaxa.5704.1.1
