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Akardita iberica n. sp. Fig. 5A ��� K 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). Etymology. The species is named after the geographic origin of the type material from the Iberian Peninsula. Other material. 1 valve, Lucena del Puerto, lower Pliocene (Della Bella coll. GDB/092); 5 valves, Lucena del Puerto, lower Pliocene (Brunetti coll. S297LDP). Diagnosis. A large species of Akardita n. gen., exceeding 40 mm in length, notably robust, with 21���24 radial ribs. 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. Distribution. The species is only known from the type locality. 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 Boll��ne, 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., Published as part of 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), pp. 215-230 in Zootaxa 4379 (2) on pages 222-223, DOI: 10.11646/zootaxa.4379.2.4, http://zenodo.org/record/1175454, {"references":["Dollfus, G. S. & Cotter, J. C. B. (1909) Mollusques tertiaires du Portugal. Le Pliocene au nord du Tage (Plaisancien). 1. Pelecypoda. Precedeed'une notice geologique. Imprimerie Nationale, Lisbonne, 103 pp.","Mayer, K. (1871) Decouverte des couches a Congeries dans le bassin du Rhone. Vierteljahresschrift der Naturforschenden Gesellschaft in Zurich, 16, 185 - 203.","Sieber, R. (1956) Die mittelmiozanen Carditidae und Cardiidae des Wiener Beckens. Mitteilungen der Geologischen Gesellschaft in Wien, 47, 183 - 234.","Schultz, O. (2003) Bivalvia neogenica (Lucinoidea - Mactroidea). In: Piller, W. E. (Ed.), Catalogus fossilium Austriae. Vol. 1. Part 2. Verlag der Osterreichischen Akademie der Wissenschaften, Wien, pp. 381 - 690.","Caze, B., Saint Martin, J. - P., Merle, D. & Saint Martin, S. (2010) Interet des motifs colores residuels des coquilles de mollusques pour la valorisation des sites paleontologiques et des collections: l'exemple du Badenien de Roumanie. In: Saint Martin, J. - P., Saint Martin S., Oaie G. & Seghedi, A. (Eds.), Le patrimoine paleontologique, Des tresors du fond des temps. GeoEcoMar, Bucarest, pp. 27 - 38.","Fontannes, F. (1882) Les mollusques pliocenes de la Vallee du Rhone et du Roussillon. Vol. 2. Acephales. Georg, Lyon, 320 pp."]}

Genus Akardita n. gen. Type species. Cardita subrevoluta de Stefani, 1888. Lower Pliocene, area of Siena, Italy. 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. 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. 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. Distribution. Lower Pliocene of the Mediterranean (northern Italy) and adjacent Atlantic Ocean (Guadalquivir Basin, southern Spain) to Recent (West Africa). 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. “ Cardita ” monodi does not fit easily in Cardita or Venericardia [cf. type species C. calyculata and V. imbricata (Gmelin, 1791) in La Perna et al. 2017, fig. 2A, B, and 3A–F]. It was recently referred to Megacardita by Huber (2010) but, as discussed by La Perna et al. (2017), this European Miocene genus includes large (up to 100 mm in length), sturdy and markedly inequilateral species. Also, its position in Cyclocardia Conrad, 1832 (Gofas & Rosenberg 2017) is questionable, as discussed below. “ Cardita ” monodi shares most shell characters with the two Pliocene species herein treated, being also particularly similar to the type species of Akardita n. gen. The specimen here studied (Fig. 1A–F), from off al- Dakhla, Western Sahara, is only slightly larger (13.3 mm in length) than the type material, which consists of two valves from distinct specimens (11.5 mm and 12.0 mm in length, respectively), referred to as holotype in the original description (Nicklès 1953: 5, pl. 1, figs 3–6). The present specimen differs by being slightly truncate posteriorly and with a lower number of ribs (17 instead of 22). In Cardita monodi, the posterior ribs are flatter and slightly narrower than the others (Fig. 1D), while they are notably finer in both fossil species, though of similar convexity. In addition, the posterior-dorsal beads are somewhat pointed and scaly (Fig. 1C, F). These sculptural differences, the only remarkable ones between C. monodi and Akardita n. gen., are not deemed to be significant. Therefore, it is assigned to the new genus as the sole living representative known so far. Akardita monodi ranges from Atlantic Morocco (35° N) to Mauritania (Baie du Lévrier, 20° N), 30–100 m depth (Ardovini & Cossignani 2004; Huber 2010, von Cosel, pers. comm. 19.10.2017). Cardita (Venericardia) matheroni Mayer, 1871, sensu Dollfus & Cotter (1909: 46, pl. 4, figs 21–26) from the Pliocene of the Tagus Valley, Portugal, may also belong in Akardita n. gen. As discussed below, it is more similar to the fossil species described herein than to Cardita matheroni Mayer, 1871, and may represent an undescribed species. Megacardita ? redoniana La Perna, Mandic & Harzhauser, 2017, from the Redonian (upper Messinian–lower Pliocene) of northwestern France, is another potential member of our new genus. La Perna et al. (2017: fig. 24A–M, 26A–G) provisionally assigned it to Megacardita, remarking its resemblance to C. monodi. Cardita zelebori Hoernes, 1865, from the lower Burdigalian of central Paratethys (La Perna et al. 2017: fig. 27A–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. 2A–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. 2E–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).

Akardita n. gen. is described for a small Pliocene to Recent group of carditids. The type species is Cardita subrevoluta de Stefani, 1888, from the lower Pliocene of Italy. The new genus includes Akardita iberica n. sp., from the lower Pliocene of southern Spain, and Cardita (Venericardia) monodi Nicklès, 1953, an extant species from West Africa. A few additional Neogene species from Europe could turn out to be representatives of the new genus, whose disappearance from European seas seems to be related to an increasing cooling trend during the Neogene–Pleistocene interval. Because of the confused status of carditid taxonomy, about which some observations are reported in the present work, it is not possible to assign the new genus to any of the traditional subfamilies.

Genus Akardita n. gen. Type species. Cardita subrevoluta de Stefani, 1888. Lower Pliocene, area of Siena, Italy. 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. 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. 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. Distribution. Lower Pliocene of the Mediterranean (northern Italy) and adjacent Atlantic Ocean (Guadalquivir Basin, southern Spain) to Recent (West Africa). 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. “ Cardita ” monodi does not fit easily in Cardita or Venericardia [cf. type species C. calyculata and V. imbricata (Gmelin, 1791) in La Perna et al. 2017, fig. 2A, B, and 3A–F]. It was recently referred to Megacardita by Huber (2010) but, as discussed by La Perna et al. (2017), this European Miocene genus includes large (up to 100 mm in length), sturdy and markedly inequilateral species. Also, its position in Cyclocardia Conrad, 1832 (Gofas & Rosenberg 2017) is questionable, as discussed below. “ Cardita ” monodi shares most shell characters with the two Pliocene species herein treated, being also particularly similar to the type species of Akardita n. gen. The specimen here studied (Fig. 1A–F), from off al- Dakhla, Western Sahara, is only slightly larger (13.3 mm in length) than the type material, which consists of two valves from distinct specimens (11.5 mm and 12.0 mm in length, respectively), referred to as holotype in the original description (Nicklès 1953: 5, pl. 1, figs 3–6). The present specimen differs by being slightly truncate posteriorly and with a lower number of ribs (17 instead of 22). In Cardita monodi, the posterior ribs are flatter and slightly narrower than the others (Fig. 1D), while they are notably finer in both fossil species, though of similar convexity. In addition, the posterior-dorsal beads are somewhat pointed and scaly (Fig. 1C, F). These sculptural differences, the only remarkable ones between C. monodi and Akardita n. gen., are not deemed to be significant. Therefore, it is assigned to the new genus as the sole living representative known so far. Akardita monodi ranges from Atlantic Morocco (35° N) to Mauritania (Baie du Lévrier, 20° N), 30–100 m depth (Ardovini & Cossignani 2004; Huber 2010, von Cosel, pers. comm. 19.10.2017). Cardita (Venericardia) matheroni Mayer, 1871, sensu Dollfus & Cotter (1909: 46, pl. 4, figs 21–26) from the Pliocene of the Tagus Valley, Portugal, may also belong in Akardita n. gen. As discussed below, it is more similar to the fossil species described herein than to Cardita matheroni Mayer, 1871, and may represent an undescribed species. Megacardita ? redoniana La Perna, Mandic & Harzhauser, 2017, from the Redonian (upper Messinian–lower Pliocene) of northwestern France, is another potential member of our new genus. La Perna et al. (2017: fig. 24A–M, 26A–G) provisionally assigned it to Megacardita, remarking its resemblance to C. monodi. Cardita zelebori Hoernes, 1865, from the lower Burdigalian of central Paratethys (La Perna et al. 2017: fig. 27A–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. 2A–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. 2E–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)., Published as part of 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), pp. 215-230 in Zootaxa 4379 (2) on pages 216-219, DOI: 10.11646/zootaxa.4379.2.4, http://zenodo.org/record/1175454, {"references":["de Stefani, C. (1888) Iconografia dei nuovi molluschi pliocenici d'intorno Siena. Bullettino della Societa Malacologica Italiana, 13, 209 - 235.","Nickles, M. (1953) Quelques lamellibranches du littoral Ouest-Africain. Institut royal des Sciences naturelles de Belgique Bulletin, 29, 1 - 11.","Dautzenberg, P. (1912) Mission GRUVEL sur la cote occidentale d'Afrique (1909 - 1910). Mollusques marins. Annales de l'Institut oceanographique, 5, 1 - 111.","Nickles, M. (1950) Mollusques testaces marins de la cote occidentale d'Afrique. Manuels Ouest-Africains. Vol. 2. Le Chevalier, Paris, 269 pp.","Pasteur-Humbert, C. (1962) Les mollusques marins testaces du Maroc. Catalogue non critique. II: Lamellibranches et Scaphopodes. Travaux de l'Institut scientifique cherifien, Serie Zoologie, 28, 1 - 184.","Bernard, P. A. (1984) Coquillage du Gabon. Shells of Gabon. Libreville, Gabon, 140 pp.","Cosel, R. von (1995) Fifty-one new species of marine bivalves from tropical West Africa. Iberus, 13, 1 - 115.","Ardovini, R. & Cossignani, T. (2004) West African seashells (including Azores, Madeira and Canary Is.). Conchiglie dell'Africa Occidentale (incluse Azzorre, Madeira e Canarie). L'Informatore Piceno, Ancona, 319 pp.","Huber, M. (2010) Compendium of bivalves. ConchBooks, Hackenheim, 901 pp.","Gofas, S. & Rosenberg, G. (2017) Cyclocardia monodi. In: MolluscaBase (2017). Accessed through: World Register of Marine Species. Available from: http: // www. marinespecies. org / aphia. php? p = taxdetails & id = 225368 (accessed 6 July 2017)","Mayer, K. (1871) Decouverte des couches a Congeries dans le bassin du Rhone. Vierteljahresschrift der Naturforschenden Gesellschaft in Zurich, 16, 185 - 203.","Dollfus, G. S. & Cotter, J. C. B. (1909) Mollusques tertiaires du Portugal. Le Pliocene au nord du Tage (Plaisancien). 1. Pelecypoda. Precedeed'une notice geologique. Imprimerie Nationale, Lisbonne, 103 pp.","La Perna, R., Mandic, O. & Harzhauser, M. (2017) Systematics and palaeobiogeography of Megacardita Sacco in the Neogene of Europe (Bivalvia, Carditidae). 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Genus Centrocardita Sacco, 1899 Type species: Chama aculeata Poli, 1795., Published as part of Oliver, P. Graham & Holmes, Anna, 2004, Cryptic bivalves with descriptions of new species from the Rodrigues lagoon, pp. 3175-3227 in Journal of Natural History 38 (23) on page 3212, DOI: 10.1080/00222930410001695123

Condylocuna io (Bartsch, 1915) Habitat. In sandy algal turf in the lower littoral. Distribution. Pointe Coton, Gravier. Remarks. See Oliver and Holmes (2004). Range. South-east Australia and South Africa. Figured specimen. 0.80 mm, NMW.Z.2000.086.00030., Published as part of Oliver, P. Graham, Holmes, Anna, Killeen, Ian, Light, Janice & Wood, Harriet, 2004, Annotated checklist of the marine Bivalvia of Rodrigues, pp. 3229-3272 in Journal of Natural History 38 (23) on page 3253, DOI: 10.1080/00222930410001695097, http://zenodo.org/record/5251926, {"references":["OLIVER, P. G. and HOLMES, A. M., 2004, Cryptic bivalves with descriptions of new species from the Rodrigues lagoon, in P. G. Oliver and A. M. Holmes (eds) The Marine Biodiversity of Rodrigues, Indian Ocean. Proceedings of the First International Marine Biodiversity Workshop for Rodrigues, 10 September to 5 October 2001, Journal of Natural History, 38, 3175 - 3227."]}

Centrocardita pileolata Oliver and Holmes, 2004 Habitat. In crevices and possibly in submarine caves, lower littoral and sublittoral. Distribution. Grand Pate´. Remarks. See Oliver and Holmes (2004). Range. Rodrigues only. Figured specimen. 2.90 mm, NMW.Z.2001.06.00039., Published as part of Oliver, P. Graham, Holmes, Anna, Killeen, Ian, Light, Janice & Wood, Harriet, 2004, Annotated checklist of the marine Bivalvia of Rodrigues, pp. 3229-3272 in Journal of Natural History 38 (23) on page 3253, DOI: 10.1080/00222930410001695097, http://zenodo.org/record/5251926, {"references":["OLIVER, P. G. and HOLMES, A. M., 2004, Cryptic bivalves with descriptions of new species from the Rodrigues lagoon, in P. G. Oliver and A. M. Holmes (eds) The Marine Biodiversity of Rodrigues, Indian Ocean. Proceedings of the First International Marine Biodiversity Workshop for Rodrigues, 10 September to 5 October 2001, Journal of Natural History, 38, 3175 - 3227."]}

Oliver, P. Graham, Holmes, Anna, Killeen, Ian, Light, Janice, Wood, Harriet (2004): Annotated checklist of the marine Bivalvia of Rodrigues. Journal of Natural History 38 (23): 3229-3272, DOI: 10.1080/00222930410001695097, URL: http://www.informaworld.com/openurl?genre=article&doi=10.1080/00222930410001695097&magic=crossref||D404A21C5BB053405B1A640AFFD44AE3

Centrocardita pileolata n. sp. (figures 124–128, 130, 131, 134) HOLOTYPE: a live collected specimen (now disarticulated and rv gold-coated), RIBW RDS 52, Grand Pate´, off Port Mathurin, 19 ‡ 39.307 ’ S, 63 ‡ 24.950 ’ E, 29 September 2001, sand in surge channel, 17 m, scuba, coll. N. L. Bruce and J. Smith, NMW.Z.2001.061.00039. PARATYPES: 2 lv (all gold-coated) and 6 rv, Grand Pate´, 18 m, NMW.Z.2001.061.00040. Measurements (mm, L 6 H). Holotype: 2.90 6 2.56. Paratypes: 4.05 6 3.65, 3.13 6 2.17, 1.60 6 1.52, 1.25 6 1.17. Description. Shell to 4 mm in length. Robust. Equivalve. Inequilateral, beaks about one-third distance from anterior. Outline trapezoidal anterior reduced; posterior dorsal margin long, straight, gently sloping to abrupt junction with truncated posterior margin, posterior ventral margin broadly rounded, ventral gently curved, anterior narrowly rounded, anterior dorsal straight and steeply sloping. Posterior area demarcated by a weak carina. Anterior and median sculpture of 13–14 strong, scaled ribs, those on the posterior carina most developed. Posterior area with a few weak riblets and a single strong rib close to the dorsal margin. Beaks prominent due to presence of a large projecting hat-like prodissoconch (figures 127, 128). Prodissoconch apex with an apical depression (figure 128) and strongly sculptured by irregular pitting (figure 127); merging into a smooth collar which is isolated from the dissoconch by a groove, in all with a diameter of 360 M m. Hinge strong. Left valve (figure 128) with two strong radiating cardinal teeth, a strong anterior lateral associated with a small ventral socket, and a denticulated dorsal area, posterior lateral longer than anterior with a deep dorsal socket. Right valve (figure 134) with a large median cardinal, anterior lateral projecting with a deep dorsal socket, posterior lateral as a marginal projection with denticulate surfaces. Inner margin deeply crenulate. External coloration white with sparse brown spots. This species broods its young and embryos were found in the holotype (figures 130, 131). Derivation of name. Latin pileolus diminutive of pileus ‘a cap’, referring to the form of the prodissoconch. Comparisons. The only other carditid found on Rodrigues is referable to Cardita variegata (Bruguière, 1792) and in its juvenile stage has a less angular outline (figures 129, 132), a simple prodissoconch and a hinge with poorly developed lateral teeth (figure 133). Viader (1951) described two small carditids from Mauritius. Cardita vaughani is equilateral with the maximum height axis running through the umbo, not behind the umbo as in C. pileolata. Cardita vinsoni is inequilateral but the ribs are nodulose rather than scaly, the prodissoconch is not capped, and the carmine and brown coloration is not present in the Rodrigues shells. Many small carditids have been described from southern Africa and southern Australia and we have reviewed the relevant literature (Bartsch, 1915; Turton, 1932; Barnard, 1964; Lamprell and Healy, 1998). Most of the smaller taxa are referred to Carditella but some to Centrocardita (s. g. Vimentum) and most are equilateral. Carditella laticosta (Smith, 1904) (South Africa) and C. (Vimentum) exulata (Smith, 1885) (South Australia) have oblique outlines but have poorly ornamented ribs., Published as part of Oliver, P. Graham & Holmes, Anna, 2004, Cryptic bivalves with descriptions of new species from the Rodrigues lagoon, pp. 3175-3227 in Journal of Natural History 38 (23) on pages 3212-3213, DOI: 10.1080/00222930410001695123, {"references":["VIADER, R., 1951, New or unrecorded shells from Mauritius and its dependencies, Mauritius Institute Bulletin, 3 (2), 127 - 155.","BARTSCH, P., 1915, Report on the Turton collection of South African marine mollusks, with additional notes on other South African shells contained in the United States National Museum, Smithsonian Institution United States National Museum Bulletin, 91, 1 - 305.","TURTON, W. H., 1932, The Marine Shells of Port Alfred, S. Africa (London: Oxford University Press).","BARNARD, K. H., 1964, Contributions to the knowledge of South African marine mollusca. Part V Lamellibranchiata, Annals of the South African Museum, 47 (3), 361 - 593.","LAMPRELL, K. and HEALY, J., 1998, Bivalves of Australia, Vol. 2 (Leiden: Backhuys Publishers), 288 pp.","SMITH, E. A., 1885, Report on the Lamellibronchiata. Reports of the Scientific Results of the voyage of HMS Challenger during the years of 1873 - 76, Zoology, 13, 1 - 341, 25 pls."]}