taxonID	type	description	language	source
7334561BFFE0FFCB1EB9FA5A7B12FF28.taxon	materials_examined	Type species. Cardita subrevoluta de Stefani, 1888. Lower Pliocene, area of Siena, Italy.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE0FFCB1EB9FA5A7B12FF28.taxon	diagnosis	Diagnosis. The new genus is uniquely characterised by its combination of the following shell characters: shape ovate-subquadrate, slightly elongate, inequilateral, weakly truncate posteriorly; length up to ca. 40 mm; approximately 20 – 25 slightly convex radial ribs, beaded in early stage, narrow and shallow interspaces, weak commarginal sculpture; hinge robust with three right and two left cardinal teeth, lateral dentition obsolete; ligament external, opisthodetic; inner shell margin crenulated, pallial line entire.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE0FFCB1EB9FA5A7B12FF28.taxon	description	Description. Shell small to moderately large, up to ca. 40 mm in length, robust, equivalve, moderately inflated. Shell shape slightly elongate, ovate, more or less subquadrate, inequilateral, well rounded anteriorly, poorly to moderately convex ventrally, slightly truncate posteriorly. Umbo relatively small, prosogyrate. Lunule small, slightly concave; escutcheon elongate, deeply sunken. Main sculpture of weakly convex radial ribs, approximately 20 to 25, with narrow, shallow interspaces. Posterior slope distinct, with weaker radial ribs. Commarginal sculpture weak, consisting of closely set, irregularly spaced growth striae crossing ribs and interspaces and producing a fine, somewhat irregular decussate pattern. Early radial ribs beaded, with slightly wider and deeper interspaces. Hinge plate moderately robust, slightly arched. Right hinge with three cardinal teeth: anterior tooth small, poorly developed; central tooth robust, triangular, elongate, strongly oblique; posterior tooth elongate, medially grooved, parallel to posterior-dorsal margin. Left hinge with two cardinal teeth: anterior tooth smaller, subtrigonal, slightly oblique; posterior tooth larger, elongate, parallel to posterior-dorsal margin. Lateral dentition only consisting of a small, tubercle-shaped anterior lateral tooth. Posterior lateral dentition almost totally obsolete. Ligament external, elongate, opisthodetic. Inner margin crenulate, especially anteriorly and ventrally. Pallial line entire, moderately convex. Anterior adductor muscle scar slightly larger, reniform; posterior scar roughly triangular; both well distinct.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE0FFCB1EB9FA5A7B12FF28.taxon	etymology	Etymology. The name was created in assonance with Cardita, type genus of the family Carditidae. Gender feminine. Included species. Cardita subrevoluta de Stefani, 1888, Akardita iberica n. sp., Cardita (Venericardia) monodi Nicklès, 1953.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE0FFCB1EB9FA5A7B12FF28.taxon	distribution	Distribution. Lower Pliocene of the Mediterranean (northern Italy) and adjacent Atlantic Ocean (Guadalquivir Basin, southern Spain) to Recent (West Africa).	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE0FFCB1EB9FA5A7B12FF28.taxon	discussion	Remarks. None of the extant European carditids, Cardita calyculata (Linnaeus, 1758), Cardites antiquatus (Linnaeus, 1758), Glans trapezia (Linnaeus, 1767), Centrocardita aculeata (Poli, 1795), Coripia corbis (Philippi, 1836), and C. jozinae (van Aartsen, 1985), mostly well-known species, shows significant similarities with both fossil species treated in the present work. Conversely, the diverse carditid fauna of West Africa (Dautzenberg 1912; Nicklès 1950, 1953 Pasteur-Humbert 1962; Bernard 1984; Cosel 1995; Ardovini & Cossignani 2004) includes Cardita (Venericardia) monodi Nicklès, 1953, which is rather similar in sculpture and shape to the two fossil species.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE0FFCB1EB9FA5A7B12FF28.taxon	description	Cardita zelebori Hoernes, 1865, from the lower Burdigalian of central Paratethys (La Perna et al. 2017: fig. 27 A – I), differs from the new genus mainly by its markedly rectangular outline with a sharper posterior truncation and a steeper posterior slope. More data on these species and better knowledge of the European Neogene carditids would provide a sounder base for their systematic position, either in the new genus or in other taxa. The only genus showing a rather close resemblance to Akardita n. gen. is Cyclocardia Conrad, 1832. According to recent authors, the genus contains about 30 living species (Huber 2010; Bouchet 2011), and several fossil species have been assigned to it (Janssen & Van der Slik 1972; Popov 1983; Janssen & Moerdijk 2004; Marquet 2005; Pérez & Del Río 2017). However, the wide morphological range encompassed by these species and an almost world-wide distribution of the group, including polar and tropical waters, suggest that too many taxa have been lumped in this genus. The type species of Cyclocardia is Cardita borealis Conrad, 1831, from the Northwest Atlantic (Canada and northern USA) (Fig. 2 A – D). Several species similar to C. borealis occur in the Atlantic and Pacific, mostly at high latitudes (Coan 1977; Coan & Valentich-Scott 2012; Huber 2010), such as the Alaskan C. crassidens (Broderip & Sowerby 1829) (Fig. 2 E – G), forming one of the few carditid groups with cold-water affinity. The easternmost record of Cyclocardia s. s. is from the Pliocene of Iceland (Vermeij 2005). Other records from the upper Cenozoic of Europe (Janssen & Van der Slik 1972; Janssen & Moerdijk 2004; Marquet 2005) are based on a few species, such as Cardita scalaris J. Sowerby, 1825, C. orbicularis J. Sowerby, 1825, and C. chamaeformis J. Sowerby, 1825, all subtriangular in shape and with finely beaded ribs, notably dissimilar from Cyclocardia s. s. Cardita scalaris is the type species of Scalaricardita Sacco, 1899, a disregarded genus which could provide a good systematic position for this group. The main differences between Akardita n. gen. and Cyclocardia s. s. concern the shell shape (Table 1). In addition, Cyclocardia s. s. has a white shell surface beneath a thick brown periostracum, while Akardita monodi has creamy to reddish blotches and chevrons on a whitish base, and its periostracum is thin and light-coloured (" cuticule mince, jaune pâle " in the original description) as could be confirmed by the examination of other material (von Cosel, pers. comm. 19.10.2017). As a component of the Northwest African fauna, Akardita n. gen. could be considered a warm-water genus. However, the latitudinal range of A. monodi is within the coastal area with strong cold upwelling (von Cosel pers. comm. 19.10.2017) that existed at least since the middle Pliocene (Vermeij 2012). It is thus warm eurythermal rather than truly tropical. Its disappearance from higher latitudes appears to be related to the mid Pliocene – Pleistocene cooling trend (Monegatti & Raffi 2001; Head & Gibbard 2005; Snyder 2016). The biogeographical history of Akardita n. gen. is similar to that of several genera whose latitudinal ranges shifted southward following the Pliocene – Pleistocene climate changes, as recently discussed for three cardiid genera (La Perna 2016, 2017; ter Poorten & La Perna 2017). The carditid genus Lazariella shows a similar shift from the Miocene Aquitaine Basin (Cossmann & Peyrot 1912) and Mediterranean (Sacco 1899) to its current occurrence along West Africa (Cosel 1995).	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE4FFC91EB9FEBE78EDFDE5.taxon	materials_examined	Type materia l. The species was described from Poggiarone, in the area of Siena. No type material has been found so far in the de Stefani and Pantanelli collections housed in the University Museums of Florence and Modena. Material examined. 38 valves, Tavernelle, lower Pliocene (Della Bella coll. GDB- 090), 35 valves (Brunetti coll., U 828 BNF 1); 3 valves, Poggio alla Fame, lower Pliocene (Brunetti coll. U 828 PGF). Illustrated material (Fig. 3 A – J) housed in MZB (47006).	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE4FFC91EB9FEBE78EDFDE5.taxon	description	Description. Shell small, slightly exceeding 20 mm in length, relatively robust, equivalve, moderately inflated. Shape slightly elongate, ovate-subquadrate, moderately inequilateral, well rounded anteriorly, poorly to moderately convex ventrally, barely truncate posteriorly. Umbo small, protruding, prosogyrate. Lunule small, slightly concave; escutcheon markedly elongate, deeply sunken. Ligament external, elongate, opisthodetic. Radial sculpture consisting of 24 – 26 poorly convex ribs. Radial interspaces narrow, shallow, poorly impressed ventrally. Growth striae distinct, closely set, crossing radial ribs and interspaces, producing a fine, irregular and decussate pattern. Commarginal sculpture mainly lamellose near ventral margin. Posterior slope distinct, with about five, notably finer ribs. Early sculpture consisting of beaded radial ribs, with slightly deeper and larger interspaces. Hinge plate moderately robust, slightly arched. Right hinge with three cardinal teeth: anterior tooth small, poorly developed; central tooth robust, triangular, elongate, strongly oblique; posterior tooth elongate, medially grooved, parallel to posterior-dorsal margin. Left hinge with two cardinal teeth: anterior tooth smaller, somewhat triangular, slightly oblique; posterior tooth larger, elongate, parallel to posterior-dorsal margin. Lateral dentition only represented by a small, tubercle-shaped anterior lateral tooth on both valves. Inner margin coarsely crenulate anteriorly and ventrally, more finely posteriorly. Pallial line entire, moderately convex. Anterior adductor muscle scar slightly larger, reniform; posterior scar roughly triangular; both well distinct.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE4FFC91EB9FEBE78EDFDE5.taxon	distribution	Distribution. Akardita subrevoluta is only known from the lower Pliocene of northern Italy.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE4FFC91EB9FEBE78EDFDE5.taxon	discussion	Remarks. The present species was first reported as Cardita revoluta Seguenza, 1877, from the Pliocene of Siena by de Stefani & Pantanelli (1878), who added a description. Ten years later, de Stefani (1888: 189, pl. 10, figs 11 – 13) noted some differences from Seguenza's species and described it as new, re-proposing the early description: " Testa cordata, subquadrata; latus anticum rotundatum, latus posticum obtuse-angulatum, margo pallearis convexiusculus: costae longitudinales 24 – 26 magnae, convexae; intervallis brevissimis separatae; transverse, praecipue ad umbones, rugosae: lunula antica parva, parum profunda: umbones recurvi obtusi: margo pallearis profunde crenatus; dens cardinalis in utraque valva brevis, in valva sinistra solidior; dentes laterales tenues, breves " (Shell heart-shaped, subquadrate; anterior margin convex, posterior margin obtusely angulate, ventral margin poorly convex: 24 – 26 radial ribs, wide, convex; separated by very narrow interspaces; commarginally rugose, especially near umbo: lunule anterior, small, slightly deep: umbos incurved, obtuse: ventral margin deeply crenulated; cardinal teeth in both valves short, stronger in left valve; lateral teeth weak, short). The valve illustrated by de Stefani (1888) (Fig. 3 K) appears more elongate and triangular than the present material, but the size reported in the description (16.5 mm in length, 16.0 mm in height) points to a more equidimensional shape, in agreement with the morphometric trend recorded in the study material (Fig. 4 A). Growth is mostly isometric, with only a slight increase in elongation with growth, as indicated by a low correlation coefficient (~ 0.5) of L / H vs L (Fig. 4 B). The maximum size is 23.2 mm in length. Width is 6.3 – 7.5 mm in 19.0 – 21.0 mm long valves.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE6FFC61EB9FA8C7D74FB80.taxon	materials_examined	Type material. Holotype, two paired valves (MZB 47004); paratype 1, RV (MZB 47005); paratype 2, RV (MZB 47005). Type locality. Lucena del Puerto, province of Huelva, Andalusia, southern Spain. Stratum typicum. Yellowish fine sands with intercalating shell beds, Arenas de Huelva Formation, lower Pliocene (Zanclean).	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE6FFC61EB9FA8C7D74FB80.taxon	etymology	Etymology. The species is named after the geographic origin of the type material from the Iberian Peninsula.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE6FFC61EB9FA8C7D74FB80.taxon	materials_examined	Other material. 1 valve, Lucena del Puerto, lower Pliocene (Della Bella coll. GDB / 092); 5 valves, Lucena del Puerto, lower Pliocene (Brunetti coll. S 297 LDP).	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE6FFC61EB9FA8C7D74FB80.taxon	diagnosis	Diagnosis. A large species of Akardita n. gen., exceeding 40 mm in length, notably robust, with 21 – 24 radial ribs.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE6FFC61EB9FA8C7D74FB80.taxon	description	Description. Shell large, up to ca. 45 mm in length, robust, equivalve, moderately inflated. Shape slightly elongate, ovate-subquadrate, moderately inequilateral, well rounded anteriorly, moderately convex ventrally, weakly truncate posteriorly. Umbo relatively small, poorly sticking out from outline, prosogyrate. Lunule small, slightly concave; escutcheon markedly elongate, deeply sunken. Ligament external, elongate, opisthodetic. Radial sculpture consisting of 21 poorly convex ribs. Radial interspaces, narrow, shallow, poorly impressed ventrally. Commarginal sculpture of closely set growth striae, producing a poorly distinct, somewhat irregular decussate pattern, ventrally replaced by coarse, lamellose sculpture. Posterior slope distinct, with 5 notably finer radial ribs. Early stage sculpture consisting of beaded radial ribs, with deeper and slightly larger interspaces than in adult stage. Hinge plate moderately robust, slightly arched. Right hinge with three cardinal teeth: anterior tooth small, poorly developed; central tooth robust, triangular-elongate, strongly oblique; posterior tooth elongate, parallel to posterior-dorsal margin. Left hinge with two cardinal teeth: anterior tooth smaller, subtrigonal, slightly oblique; posterior tooth larger, elongate, parallel to posterior-dorsal margin. Lateral dentition represented, on both valves, by a small, tubercle-shaped anterior lateral tooth, and by a shallow, elongate, barely seen posterior thickening, close to hinge plate ending. Inner margin coarsely crenulated anteriorly and ventrally, posteriorly almost smooth. Pallial line entire, moderately convex. Anterior adductor muscle scar slightly larger, reniform; posterior scar roughly triangular; both well distinct. Measurements. Holotype, L 42.5 mm, H 37.1 mm, W 13.7 mm; paratype 1, L 42.6 mm, H 38.6 mm, W 15.4 mm; paratype 2, L ~ 40 mm (estimated), H 39.5 mm, W 13.1 mm. Other material up to L 45.9 mm.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE6FFC61EB9FA8C7D74FB80.taxon	distribution	Distribution. The species is only known from the type locality.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
7334561BFFE6FFC61EB9FA8C7D74FB80.taxon	discussion	Remarks. The available material shows a moderate variability in rib number, ranging from 21 to 24. The new species differs from A. subrevoluta by being much larger (up to ca. 45 mm in length, while A. subrevoluta does not exceed 25 mm), markedly more robust, slightly more inequilateral, ventrally more convex, umbo less prominent, and rib number with a limited overlap at 24 ribs. The new species resembles the undescribed species from the Pliocene of Portugal, here referred to as Cardita matheroni sensu Dollfus & Cotter, 1909. According to illustrations by Dollfus & Cotter (1909: pl. 4, figs 21 – 26), the species from Portugal has poorly convex ribs without nodulose sculpture. It differs from Akardita iberica n. sp. by being more rounded in outline and by the occurrence of some particularly deep, irregularly spaced growth striae, in addition to the finer ones. Both species are similar in size, exceeding 40 mm in length. The original Cardita matheroni Mayer, 1871, is from the upper Miocene Congeria beds in the area of Bollène, southern France (Mayer 1871). According to Mayer, his species is similar to Cardites antiquatus and to Cardita partschi Goldfuss, 1840. The Miocene C. partschi is in turn similar to C. antiquatus, with nodulose ribs (Sieber 1956; Schultz 2003; Caze et al. 2010). Cardita matheroni sensu Fontannes, 1882, from the Pliocene of the Rhone Valley (Fontannes 1882: 122, pl. 7, fig 15 – 17) seems to be yet another distinct species.	en	Perna, Rafael La, Brunetti, Mauro M., Bella, Giano Della (2018): Systematic position of two Pliocene carditids with description of Akardita n. gen. and A. iberica n. sp. (Bivalvia: Carditidae). Zootaxa 4379 (2): 215-230, DOI: 10.11646/zootaxa.4379.2.4
