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Subfamily unassigned Maoricardium pseudolatum Voskuil& Onverwagt, 1991 Maoricardium setosum (Redfield, 1846) Nemocardium fulvum ter Poorten, 2013

Subfamily Cardiinae Vepricardium asiaticum (Bruguière, 1789)

Subfamily Fraginae Lunulicardia retusa (Linnaeus, 1767)

Subfamily Laevicardiinae Fulvia australis (G.B. Sowerby II, 1834) Lyrocardium lyratum (G.B. Sowerby II, 1840)

Subfamily Orthocardiinae Frenexicardia victor (Angas,1872)

Subfamily Trachycardiinae Acrosterigma attenuatum (G. B. Sowerby II, 1841) Acrosterigma biradiatum (Bruguière, 1789) Acrosterigma impolitum (G. B. Sowerby II, 1834) Acrosterigma maculosum (W. Wood, 1815) Acrosterigma oxygonum (G. B. Sowerby II, 1834) Vasticardium angulatum (Lamarck, 1819) Vasticardium assimile (Reeve, 1844) Vasticardium assimile lacunosum (Reeve, 1845) Vasticardium elongatum cipangense (Vidal, 1993) Vasticardium indioceanum (Vidal, 1993) Vasticardium pectiniforme (Born, 1780) Vasticardium rubicundum (Reeve, 1844) Vasticardium subrugosum (G.B. Sowerby II, 1839)

Subfamily Tridacninae Tridacna maxima (Röding, 1798)

Fulvia asiatica (Bruguière) Vasticardium arenicola (Reeve) - Kirtisinghe, 1978 Names unverified, identities unknown

PLATE 22

COCKLES Family: CARDIIDAE

Reported by other workers and not included in this catalogue

The family Cardiidae is very large, with numerous genera. The shells vary in size from small/medium to large, thin-shelled and light to thick-shelled and heavy, glossy to strongly sculptured, and of various shapes. They are distributed in a number of subfamilies, based on the external appearance and other characters. They are presented here grouped in subfamilies, and not according to the alphabetical order of the generic name as in other family plates, as an aid to the identification of a specimen in the hand. There have been many changes in the generic names in recent years.

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Subfamily Unassigned

MF238: ?Mount Lavinia, Bellangala area. MF239: Dehiwela, Auburnside reef N, 3

m, empty in cavity. MF579: Kayankerni, Thennadi Bay, beached.Also, beached at Erakkandy, Trinco.

Maoricardium pseudolatum Both species listed in Checklist 2012 and Kirtisinghe, 1978 under the generic name Afrocardium. The tapering rounded posteriors are longer than in M. setosum. Flat ribs with tubercles in rows down the midline. Creamy white with cloudy concentric bands of orange under abrown periostracum with long, stiff, bristle-like hairs attached to midline tubercles. Hairs fragile when dried. Labels destroyed by insect attack, many collections from Bellangala, Auburnside Godagala, Mount Lavinia beach between 1978 & 1980. Live collections byssally attached in rocky crevises or under sponges. Shells have a narrow posterior byssal gape.

MF236: No collection data. Periostracum missing. Differs from M. pseudolatum in the shorter, blunt posterior.

Nemocardium fulvum A fragment of a single valve found only once, in July, 2019. The definitive identification established in March, 2020 with reference to the paper by ter Poorten, 2013. “Type locality India, S.E. part, Tamil Nadu, off Colachel, trawled by fishermen, depth 70-125 m” - WoRMS. The genus is said to occupy deep water, growing to 60-80 mm. This species widely distributed over the Indo-Pacifc from Madagascar to the north of Australia. Poorten, J. J. ter. (2013). Revision of Recent species of the genus Nemocardium Meek, 1876 (Bivalvia, Cardiidae), with the descriptions of three new species. Basteria. 77 (4-6): 45-73.

MF823, Maggona, Thudawa Small Boat Fishery Harbour, fishing trash. Single r/v fragment. Anterior and ventral margins convex, the posterior truncate. Characteristic sculpture of spinose radial ribs posterior 1/3, faint cancellate markings rest of shell, lunule quite smooth.

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Subfamily Cardiinae

MF280: Kalpitiya, Kandakuliya, beached, single valve, 32 ribs. MF450: Kalpitiya, Puttalam Lagoon west shore,fishing trash.

Also, Mannar Island, Pesalai anchorage, fishing trash;Silavaturai, Doric beach, beached; Wellawatte, seaward of Palagala reef, mud/sand bottom, snagged on crab net.

Vepricardium asiaticum In Vepricardium length = height, in Trachycardium, length is less than the height. 32 ribs with a heart-shaped lunule. Anterior ribs narrow, wider rounded interspaces. Medial ribs round-topped, equal-sized flat interspaces. Posterior ribs skewed with lamella extensions. Ribs covered with tiny elongated spines on sides and rounded tubercles above, giving a fuzzy appearance.("with a series of erect lamellate spines forming a prominent, continuous palisade near shell margin" - Poutiers, 1988.) Interior markedly ribbed over umbonal region.

Subfamily Fraginae

Lunulicardia retusa Checklist 2012 as Fragum hemicardium; and Kirtisinghe, 1978 p.29 as Lunulicardia subretusa, another synonym. Described from single valves. Shell distorted with posterior compressed and flattened, set off from the anterior by a strong keel. Umbones strongly incurved; lunule strongly depressed forming a cavity. Posterior margin gently convex, meeting the antero-ventral at an acute angle. Antero-dorsal meets antero-ventral at a well rounded junction. Two cardinal and two lateral teeth, the posterior laterals closer to the cardinals. Interior ribbed, margin crenulate corresponding to the external sculpture. Sculpture of low radial ribs, stronger and wider over the anterior, the narrow grooves between pitted and crossed by bars that also cross many of the ribs. Lunulicardia is distinguished from the similar genus Hemicardium by the strongly depressed lunule.

MF525: Wattala, Pegasus Reef hotel beach, beached, numerous weathered single valves; also, MF691: Unawatuna, in-shore rock fringe, <1m, sand, weathered s/valves; MF760: Erakkandy estuary mouth, weathered s/v.

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Subfamily Laevicardiinae

(Left) MF275: Trincomalee, Kinniya, empty, by diving. 45 ribs. (Centre) MF272: Trincomalee, Marble Bay, 2m, empty, by diving. (Right )MF273: Negombo, 15m, alive, buried in sand, by diving. 53 ribs. Also, MF274: Clappenberg Bay, empty, beached, 20 x 21 x 19mm. 46 ribs. MF795: Tincomalee harbour, Gt. Sober Island, 3m, empty, muddy sand bottom.

Fulvia australis Shell lightweight, inflated, transversely oval with drawnout posterior, umbo small. Ribs numerous, generally flattened and smooth, delineated by incised radial lines except anteriorly where they are finely granulated and posteriorly where they are slightly elevated and crossed by growth lines. Whitish, umbonal region purplish-pink. Interior white, umbonal and posterior blushed with pink. Beaks brownish pink. Listed in the 2012 checklist as species ofLaevicardium. Small difference in shape and colouration, seen in the three images above, made identification difficult. The final determination as F. australiswas based on online images and descriptions in articles by ter Poorten and in the idscaro website.

(Left) MF192: Colombo, Vatiya Parai (reef), 28 m, sand bottom, empty, by diving. (Centre) MF461: Maggona, Thudawa Bay, fishing trash. (Right) MF191: Colombo, Degal Meda (reef), 24 m, sand, empty. Also, MF190: Beruwela, beach, single l/v 36.4x40 mm.

Lyrocardium lyratum Lyrate cockle Checklist 2012 as Nemocardium lyratum; Discors lyratum is another synonym. Solid, swollen and sub-spherical; Ht > L. Strong radial riblets posteriorly, oblique threads over the anterior half. Periostracum pink-red marbled with white over a white shell. Interior white blushed with pink in the umbonal region. Prominent teeth.

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Subfamily Orthocardiinae

Frenexicardia victor Victor's prickly cockle Inflated, solid, somewhat globular shell with spinose ribs over the whole surface. The anterior margin convex, the posterior angulated. Ornamented with spaced ribs bearing recurved pointed spines, the points directed towards the umbo. Creamy white with blotches of deep rose pink that includes the spines within the coloured patches. Interior apricot, the crenulated margin ground colour. De Bruyne, 2003 gives depth range as 25-200 m, rarity as 'uncommon to relatively rare'. This has been found only from Maggona.

(Left) MF551; (Right) MF741. Three collections, all from Maggona, Thudawa bay Small Boat Fishery Harbour, fishing trash.

Subfamily Trachycardiinae

Acrosterigma: The shells are pear-shaped with pointed umbones. Six species are included here, the sculpture ranging from barely perceptible ribs to strong ribs.

MF622: Wattala, Pegasus Reef Hotel & Dikkowita beaches, beached single valves. Numerous. Mostly much weathered, damaged and rust-stained. Type locality Ceylon (Sri Lanka).

Checklist 2012 as Laevicardium biradiatum, a synonym. MF144: Beruwela, shoal, 26m, empty, by diving. Shell thin, glossy. Numerous low transversely ridged riblets. Type locality Sri Lanka.

From Jaffna, Mandaitivu, fishing trash, empty. 36 rounded ribs crossed by low transverse bars the posterior 6-7 ribs acute with granules. Periostracum thin, brownish.

Acrosterigma maculosum Checklist 2012 as Laevicardium sp. 2. MF271: No collection data(Trincomalee, Kinniya?), one specimen. The two pink internal radial streaks in the umbonal region are characteristic. 69 ribs. Lightweight, inflated, small pointed umbo, widening to a wide rounded ventral margin.

Ribs numerous: narrow, rounded, transversely ridged, 7 post ones flat, tuberculated; interspaces narrower, rounded. Cream coloured with purplish blotches, beaks pink. Interior white, surface colours showing through, two pink streaks fanning out from beaks.

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MF285: Colombo, Palagala (reef), inshore, 13 m, sand, single valves, 38 ribs.

L/v 30x33 mm

From unknown location, beached single valve. Also collected beached single valves from Wattala sites. 38 to 41 ribs.

MF620: Wattala, Dikkowita beach, single valve much rust-stained. 36.65 x 44.25 mm, 38 ribs.

Acrosterigma oxygonum Shells taller than long. Ribs 38 to 41, keeled, triangular in section, interspaces narrow. Crossed by threads passing over the summits and both sides but not over the interspaces. The 9 posterior ribs flat with tubercles. Interior radially grooved corresponding to the external ribs, margin crenate. All specimens much weathered, the Wattala ones often covered with tar, rust stained or discoloured. Abundant on the Wattala beaches. Photo of one of the few showing good colour (at left): white with pink-red clouds. Similar colour seen faintly in one of the two Colombo specimens.

MF277: Batticaloa, Kalmunai,14 m, empty on sand, by diving. Gaping of anterior and posterior characteristic. Marginal view shows rounded ribs with wide interspaces. 32 ribs.

MF278: Trincomalee, Nilaveli, Pigeon Is., Salabalas, 9 m, empty, by diving. 31 ribs.

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Vasticardium angulatum Checklist 2012 as Trachycardium angulatum. Large, heavy, elongated oval shells with small umbos and rather flattened anterior and posterior margins. 30-34 wide, rounded ribs, flat interspaces about half as wide. Ribs crossed by transverse ridges: across whole width anterior, shortened to oblique linear tubercles from the middle onwards; the posterior 7-8 with a row of radially aligned spine-like tubercles. Concentric cloudy bands of orange-pink or tan and white, covered by a brown periostracum. Interior white with the crenulated margin gold, more extensive posteriorly. Some golden blushing in the umbonal region. "Salmon and beige coloured forms occur along the East coast of Africa..." (ter Poorten, 2007). Also, MF298: Off Colombo, Gigiripita shoal, 21.5 m, empty on sand, by diving. 50.5x60x37 mm, tan and white marbling, brown periostracum.

MF599: Jaffna, Mandaitivu, fishing trash. A highly coloured single valve. 30 ribs.

MF671: Trincomalee, Erakkandy lagoon mouth, beached. 51.18 x 64.36 x 45.36mm. 31/32 ribs.

MF696: Jaffna, Kachchativu Is., beached. Coll. Devaka Weerakoon & Arjan Rajasuriya.

MF282: Trincomalee; Dutch Bay, empty, 281 Marble Bay, 2 m, on sand amongst rocks, 283 Nilaveli, Pigeon Islands, off Big Is. beach, 4 m, empty, amongst corals.

Vasticardium assimile Shell somewhat rectangular, inflated. Anterior margin gently convex, posterior margin flattened, ventral margin obliquely convex. Ribs 31/32, anterior ones broadly tabulate with narrower, rounded interspaces, becoming narrow tabulate with 'V'-shaped interspaces behind the midline, the most posterior 7/8 low, narrow, with erect nodular spines; other ribs surmounted by transverse bars over the summits and some way down the sides but not across the interspaces. Interior white, faintly grooved corresponding to the external ribs, margin crenulate.

Vasticardium assimile lacunosum Checklist 2012 as Vasticardium lacunosum Rather squarish shells, moderately inflated. 30-33 wide, rounded ribs; flat interspaces narrow. Ribs transversely ridged, sometimes lacking over the midline at the centre but extending down the sides and across the interspaces. Posterior ribs 7 or so, scaly. Colour yellowish grey or cream, some brownish with varying degrees of blackish, grey or brown staining on the ribs arranged in concentric lines over the posterior half. Interior porcelain white.

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Vasticardium elongatum cipangense The collections previously determined as Trachycardium enode (checklist 2012) is deemed a mis-identification as V. enode is a Japonic species. (See NMR Picture pages for an image of a shell from Rameswaran, India.) Three single valves are in the collection, all three from Colombo, found on the sand bottoms adjacent to the reefs of Palagala and Degal Meda. These shells are distinguished by their compact, squarish outline with close-set, angular ribs as against the oval shape of V. indioceanum. The exterior colour is light tan with white clouding under a shiny brown periostracum, the interior white with a yellow umbonal patch.

MF290: Colombo, Degal Meda (reef), 24 m, single right valve, by diving. 43 ribs.

MF291: Colombo, Palagala (reef), 14 m, single right valve, by diving. 40 ribs.

MF279: No collection data, elongated oval, angular ribs, 43 ribs.

MF785: Jaffna, Mandaitivu. Single l/v 69 x 90mm, 39 ribs, periostracum intact, brown, over creamy white shell with pink marbling over umbo. Interior white, marginal pink posteriorly.

Vasticardium indioceanum (Checklist 2012 as Trachycardium elongatum) Elongate cockle Vasticardium elongatum was described as occurring as six subspecies throughout the tropical Indo-West Pacific up to the Tonga Islands. This position was challenged and the subspecies have been subsequently raised to specific rank. V elongatum elongatum is now V. indioceanum.

Shells oblong-ovate, large and heavy with 39 - 43 tabulate ribs with narrow rounded interspaces.

MF596: Trincomalee, Great Sober Island, 3m, empty, muddy-sand, heavily encrusted. 29 - 31 ribs.

Vasticardium pectiniforme Pacific yellow cockle Shells solid, higher than long. 29-31 rounded ribs crossed by arcuate ridges interspaces narrow the posterior 7 scaled. Margin strongly crenulate inside. Cream coloured, interior white.

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MF289: Kalpitiya, Bar Reef. Empty, by diving. Intact periostracum.

MF580: Kayankerni, Thennadi Bay. Beached single valve, denuded.

Granular ornamentation anterior, transverse bars on tabular ribs medial, rounded spines on narrow ribs posterior.

Vasticardium rubicundum Small broadly oval shells with slightly flattened anterior and posterior margins, ventral rounded. Inflated. Ribs 32 - 34, narrow, tabulate; inter-spaces as wide, round-bottomed. Ribs transversely ridged throughout anterior and medial; 6-7 posterior ribs with thick scales. With periostracum colour tan or hazel, interspaces creamy, chestnut streaks scattered on ribs in groups forming blotches. Umbo pale. Interior white, the crenulated margin chestnut. Denuded, colour cream with purplish markings, interior margin purple. Umbonal region with two radial streaks of purple with a variable diffuse pink flare; may be absent. Also, MF288, Batticaloa, Kalmunai, beached s/v; MF287, off Ratmalana, Deep Reef, 19 m, empty on sand at edge of reef amongst rocks.

Vasticardium subrugosum Checklist 2012 as Vasticardium flavum. V. flavum subrugosum (with strongly developed rough ribs) accepted as V. subrugosum. Shell nearly circular, small umbo, not very inflated. Ribs 31, round-topped, vertical-sided; narrower interspaces flat, transversely ridged - the anterior well- spaced, medial closely spaced; posterior with erect scales. White, interior white with a light yellowish staining postero-ventral edge.

MF286: Trincomalee, Dutch Bay, 3m, empty on sand amongst rocks, 31 ribs by diving.

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Subfamily Tridacninae

Tridacna maxima The large byssal opening is behind the umbones, the long ligament is in front. In life the shell is attached with the ventral margin uppermost. Shells are pictured in the living position. The single posterior muscle scar merges with that of the large posterior pedal retractor that is smaller. There is no pallial sinus. Another species may be in the area—T. squamosa is said to grow larger, fan-shaped, the grooves between the radial folds as wide, the erect scales larger and more widely spaced. (Poutiers, 1998 p. 271)

MF015: From Colombo in-shore reefs, by diving.

MF014: Trincomalee, Coral sands bay, 2-3m, by diving.

Batticaloa, Kattankudy, 1905 Batticaloa, 1903 Kirinda, Great Basses reef, 1903

Three under water pictures show the shells with the mantle partially retracted. As they are sensitive to light intensity, at the approach of a diver the mantle begins to retract and the shells to start to close. When fully open the scales are completely covered by the mantle lobes.

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Bibliography

Citation: ter Poorten, J. (2014). Cardiidae. In: MolluscaBase (2016). Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=229 on 2017-01-10.

Citation: ter Poorten, J. (2014). Acrosterigma attenuatum (G. B. Sowerby II, 1841). In: MolluscaBase (2017). Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=345700 on 2017-04-06

De Bruyne, R. H. (2003). The Complete Encyclopedia of Shells, Rebo Productions, Lisse, The Netherlands.

Images of Fulvia fragilis and Fulvia australis compared available at: http://www.idscaro.net/sci/04_med/class/fam5/species/fulvia_fragilis1.htm

Kirtisinghe, Parakrama (1978). Sea shells of Sri Lanka, Tuttle, Tokyo.

Poutiers, J. M. (1998). Bivalves (Acephala, Lamellibranchia, Pelecypoda). In: Carpenter, K. E. and Niem, V. H. (eds.), FAO Species Identification Guide for Fishery Purposes, The Living Marine Resources of the Western Central Pacific, Vol. 1, FAO, Rome.

Reeve L.A. (1844-1845). Monograph of the genus Cardium. In: Conchologia Iconica, vol. 2, pl. 1-22 and unpaginated text. L. Reeve & Co., London. [stated dates: pl. 1-4 [3 undated], October 1844, pl. 5-8, November 1844; pl. 9-12, December 1844; pl. 13-16, January, 1845; pl. 17-22, March 1845], available online at: http://biodiversitylibrary.org/page/8937393.

ter Poorten, J. (2007). Results of the Rumphius Biohistorical Expedition to Ambon (1990) Part 13. Mollusca, Bivalvia, Cardiidae, Zoologische Mededelingen, 81-1 (June 2007), available online at http://www.zoologischemededelingen.nl/81/nr01/a15

ter Poorten, J. (2012). Fulvia (Fulvia) nienkeae spec. nov., a new Fulvia from the Central Indo-West Pacific (Bivalvia, Cardiidae), Basteria76 (4-6): 117-125 (2012),available online athttps://www.researchgate.net/publication/258374844.

Poorten, J. J. ter. (2013). Revision of Recent species of the genus Nemocardium Meek, 1876 (Bivalvia, Cardiidae), with the descriptions of three new species. Basteria. 77 (4-6): 45-73.

Vidal, J., 1997. Taxonomic revision of the Indo-Pacific Vasticardium flavum species group. - Zoosystema 19(2-3): 233-253. (http://www.zoologischemededelingen.nl/81/nr01/a15)

Acrosterigma attenuatum original description: Sowerby II, G. B. (1841). [An extensive series of new species of the genus Cardium was exhibited by Mr Cuming ...]. Proceedings of the Zoological Society of London 8 (92): 105-111 (May 1841); Annals and Magazine of Natural History, ser. 1, 7 (46): 506-511 (1 August 1841). [Type locality Labelled ‘Ceylon, Zanzibar, Philippines, M.C.’ although only Ceylon [Sri Lanka] is mentioned in the original description. Coll. Cuming.] WoRMS.

Acrosterigma biradiatum: Type locality Sri Lanka. Vidal J. 1999. - Taxonomic review of the elongated cockles: genera Trachycardium, Vasticardium and Acrosterigma (Mollusca, Cardiidae). Zoosystema 21(2): 259-335.

Acrosterigma impolitum: Original description Sowerby, G. B. I & Sowerby, G. B. II. (1832-1841). The conchological illustrations or, Coloured figures of all the hitherto unfigured recent shells. London, privately published., available online at http://www.biodiversitylibrary.org/item/109863.

Vasticardium indioceanum: Original description (of Acrosterigma elongatum indioceanum Vidal, 1993) Vidal J. (1993) Variability of Acrosterigma elongatum, a polytypic species (Mollusca, Cardiidae). Journal of the Malacological Society of Australia 14: 41-58. [30 October1993] page(s): 53.

MolluscaBase eds. (2020). MolluscaBase. Vasticardium indioceanum (Vidal, 1993). Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=1048886 on 2020-04-01.

MolluscaBase eds. (2020). MolluscaBase. Vasticardium elongatum cipangense (Vidal, 1993). Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=381326 on 2020-04-02

Poorten, J.J. ter, 2005. Outline of a systematic index - Recent Cardiidae (Lamarck, 1809). VISAYA net. (Updated 2009 for

WoRMS), available online at http://www.conchology.be/en/shelltopics/visaya-net/date.php?year=2005

MF: 10.1.2017; 3.2.2017; 27.4.2017; 1.5.2017; 3.4.2020