Arcuatula capensis (Krauss, 1848) Botula cinnamomea (Gmelin, 1791) Brachidontes pharaonis (P. Fischer, 1870) Brachidontes variabilis (Krauss, 1848) Ciboticola lunata (Hedley, 1902) Gregariella coralliophaga (Gmelin, 1791) Jolya elongata (Swainson, 1821) - subfossil Leiosolenus lima (Jousseaume in Lamy, 1919) Leiosolenus malaccanus (Reeve, 1857) Leiosolenus obesus (Philippi, 1847) Lithophaga teres (Philippi, 1846)

Modiolus auriculatus (Krauss, 1848) Modiolus modulaides (Röding, 1798) Modiolus philippinarum Hanley, 1843 Musculus cupreus (Gould, 1861) Mytilus unguiculatus Valenciennes, 1858 Perna perna (Linnaeus, 1758) Perna viridis (Linnaeus, 1758) Septifer bilocularis Linnaeus, 1758

Lithophaga nigra (d'Orbigny, 1853)

- Kirtisinghe, 1978

Reported by other workers and not included in this catalogue

Ligament antero-dorsal in Mytilids Anterior partition in Septifer Teeth-like crenulations behind ligament in

Brachidontes

PLATE 1

MUSSELS, CORAL BORERS ETC Family: MYTILIDAE

The shell valves are equal, elongated or oblique, generally triangular in shape with a pointed anterior and an

expanded, rounded posterior; or cylindrical, usually with a narrow byssal gape at the ventral margin. The umbones are

at or near the anterior end of the shell. Externally smooth or radially ribbed, the ribs when present being stronger on the

posterodorsal and anterior areas, reduced on the ventral medial area. Periostracum present, being smooth, flaky or

hairy. The ligament is external, deeply set and supported by a whitish ridge, along the anterior dorsal margin. Hinge

teeth are absent or reduced; small marginal crenulations sometimes present behind the ligament (in Brachidontes

Smith, 1964). The anterior adductor muscle scar small or absent, the posterior scar large. The pallial line without a sinus.

Internal margins smooth or crenulate.

Mussels are sedentary animals, attached to hard substrates by a byssus, sometimes nestlers, or boring into coral

or rock, or associated with ascidians. The boring forms have special acid-secreting glands in their mantles. Found in all

regions from the intertidal zones down to great depths. Some species are edible. Many species are farmed.

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Arcuatula capensis

Arcuatula capensis Estuarine mussel Checklist 2012 as Brachiodontes sp. 2 - (sic)

This genus resembles Brachidontes but does not have the dorsal marginal crenulations behind the ligament. Is a shallow brackish water inhabitant.

Elongated slender shells, somewhat inflated, anterior obtuse, posterior drawn out, dorsal elevation rounded, ventral concave. Beaks behind anterior end. Nodular riblets divaricate from beaks to dorsal, posterior and ventral margins as well as a recurved set forward to the anterior extremity. Periostracum dusky brown, dull, striped pattern showing through. Shell glossy, whitish with a "turkey wing" pattern of cinnamon wavy, concentric lines tending to smudge behind making that portion darker. This species differs from the marine species of Brachidontes in being slenderer with a drawn out posterior and a protuberant anterior, beaks behind the anterior extremity and riblets curving forwards into it. Is also attractively patterned underneath a dull dusky brown periostracum.

MF356: Hendala, Hamilton canal, byssally attached on rocks of embankment.

MF438: Chilaw, Mundal lake, empty, on dead Halophila and oysters. 10: 16-11 mm; 3: 7-5 mm. Marooned on shore by receding water. Juveniles on Magallana bilineata below and on exposed mangrove air roots above.

Botula cinnamomea

MF554: Maggona: Thudawa Bay, SBFH. Boring into coral. These are chemical borers found in tight fitting cavities.

MF563: Maggona, Thudawa Bay, SBFH, fishing trash. In coral bore-hole, with periostracum, no external opening.

MF742: Maggona, Thudawa Bay, SBFH, fishing trash. In sandstone bore-hole, with periostracum.

Botula cinnamomea Cinnamon coral borer Checklist 2012 as Lithophaga cinnamomea (Kirtisinghe, 1978 p. 14). Shell rectangular, not gaping, beaks incurved, protruding forwards. External ligament set in a serrated groove. Exterior with growth lines only. No pallial sinus. Light tan colour. Periostracum glossy, dark brown. Up to 25mm in length. A shell boring into coral, occupying tight fitting cavities that it creates by chemical means. The specimens found had all been sequestered within their chambers by overgrowth of the coral.

- Valentich-Scott& Tongkerd, 2008 Fig. 2 A-E. See also Valentich-Scott & Dinesen, 2004 discussion under B. fusca.

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Brachidontes

Brachidontes pharaonis Batticaloa ribbed mussel Checklist 2012 as Brachiodontes sp. 3 (sic) Shells stubby, wedge-shaped, moderately inflated, the beaks very close to the anterior extremity. A series of crenulations characteristic of the genus lie just behind the ligament. Strong, nodular radial riblets diverge from the beaks, being weak ventrally. The shell is coloured reddish-brown and overlaid with a thick hairy periostracum. Up to 13.5 mm in length. A shallow-water marine species. Distributed Indian Ocean and spread into the Mediterranean Sea.

- The Mediterranean Science Commission website

MF517: Batticaloa: Palameenmadu, lagoon. Periostracum denuded (above); with hairy periostracum (below). Inset in r/h image: detail of post-ligament crenulations. Byssally attached to clumps of oysters and crevices on the undersides of rocks lying on the muddy sand bottom in very shallow water.

Brachidontes variabilis Common ribbed mussel Checklist 2012 as Brachiodontes sp. 1(sic).

Shells small, elliptical in shape, inflated. The beaks level with the anterior extremity giving a smoothly rounded outline. Fine nodular riblets divaricate from beaks to dorsal, posterior and ventral margins. The periostracum is strong and glossy, the shell coloured a rich chocolate brown. Up to 12.7mm in length. An intertidal marine species. Also, Wattala, Dikkowita, beached; Beruwela, Barberyn Reef, rock pools, amongst algae.

- Smith, 1964 p. 151.

MF355: Hendala, beach rock platform. Byssally attached under tufts of algae on intertidal rocky shore. From 7 mm to 12.65 mm. Located by Suki Ekaratne.

MF351: Mount Lavinia, Dig Gala. Byssally attached on wave-washed rock at edge of beach, overgrown by algal tufts. Inset shows post ligament crenulations.

Ciboticola lunata Checklist 2012 as Septifer sp. 1 Shell laterally compressed, inflated ventrally with a ventral surface that makes a right angle with the sides. Elliptical in shape with a pointed beak turned down slightly; antero-dorsal hinge margin straightish, shorter than the postero-dorsal margin that is smoothly rounded ending in a narrowly rounded postero-ventral angle. The ventral margin is arcuate with a gap for a byssus. A sharply angled keel from the beak to the postero-ventral angle separates the gently rounded side of the shell with the flattened, somewhat excavated, narrow

MF466: Colombo, Wellawatte, beyond Palagala (reef). Fishing trash from bottom set crab nets on muddy-sand bottom. The right valve has been denuded.

ventral surface. Internal shelly septum anteriorly as in Septifer. The shell is white with some reddish-brown staining along the hinge margin and covered with a reddish-brown hairy periostracum. The hairs are quite coarse and long over the posterior parts. Described from Australia.

- Images in WoRMS; biolib.cz; WMSDB

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Gregariella coralliophaga

Gregariella coralliophaga Pointed borer Shell small, trigonal, with straight dorsal margin, anterior umbones above a rounded anterior extremity, sinuate ventral margin and pointed posterior. Edentulous, no obvious ligament attachment. A keel from umbo to acute postero-ventral angle defining a broad, concave, postero-dorsal slope that is at right angles to the medial aspect. A shallow radial groove runs from umbo to ventral margin that is notched. No gape, no byssal notch. Sculpture of crowded radial threads in front of the radial groove, smooth behind. Postero-ventral slope crossed by coarser granulose radial threads. Margins crenulate corresponding to external sculpture, absent adjacent to smooth area. Anterior adductor scar circular, posterior linear, pallial line entire. Shell white, periostracum yellowish-brown with hairs developed behind. 11 to 13mm in length. A coral-associated species, probably more a nestler than a borer.

- Ranong, Thailand - "A small, hairy mussel found in crevices on intertidal rocky shores." MF675: Maggona, Thudawa Bay, SBFH fishing trash. Inside cavity in coral rubble covered by calcareous concretion. MF676: Trincomalee, Erakkandy lagoon mouth beach. Inside cavity in coral rubble. Periostracum removed (bottom). In cavities of dead coral rubble, fishing trash or beached.

Jolya elongata

Jolya elongata A single mineralised shell from the Negombo 'Fossil Reef' where many species of similarly mineralised bivalves, gastropods and echinoderm tests have been found. (See article “Beach-combing surprises - Fossils or Subfossils?” available on this website).

Anterior view Dorsal angle rounded, far behind midline, postero-dorsal margin descends at 45` to a narrow, rounded posterior pointing downwards. Beaks behind blunt, high anterior margin. Shell not very inflated the anterior part sub-cylindrical. - Dance, 1977 (description); many images on-line, from Thailand and the Philippines.

MF189: Negombo, ‘Fossil reef’, 14 m, mineralized. Sand bottom together with other subfossil shells, by diving.

Leiosolenus Boring Date Mussels of the genus Leisolenus develop extra-periostracal concretions

Leiosolenus lima Crusted date mussel Dorsal margin angulated, antero-dorsal and postero-dorsal margins sloping; beaks sub-terminal with gap between beaks in dorsal view; tapering both anteriorly and posteriorly to rounded ends, ventral margin convex. Concretions thick, sculptured: finely granular over anterior, ventral and dorsal aspects becoming coarse postero-ventrally, a wedge-shaped area extending from a mid-dorsal point to include entire posterior with strong erect, divaricating laminae, hardly extending beyond posterior of shell.

- Valentich-Scott& Tongkerd, 2008 p. 196 Fig. 5 A-E.

Maggona, Thudawa Bay, fishing trash, boring into sandstone. MF700: Left - Denuded shell below. MF686: Right. Both images show the angulated dorsum and the characteristic extra-periostracal concretions that do not extend beyond the end of the shell.

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Leiosolenus malaccanus

Left - MF312: Colombo, Kelani Gala, 17 m, boring into a Chama sp. Centre - MF314: Mount Lavinia, Bellangala, inshore, 3 m, boring into underside of aFavia colony. Right - MF689: From a pooled collection from Colombo & Trincomalee, boring into corals. Leisolenus malaccanus Malacca date mussel Dorsal angulation low; beaks dorsal, sub-terminal, no gap between right and left; antero-dorsal sub-parallel to ventral, anterior profile blunt, broadly rounded; postero-dorsal sloping, rear tapers to narrow rounded posterior. Thinly covered with concretions, not sculptured, conical extension beyond posterior of shell following shell contour.

- ranong.myspecies -- " A borer into coral and limestone. A distinctive, pitted, calcareous deposit extends beyond the posterior margin of the shell."

- Valentich-Scott & Tongkerd, 2008 p. 197 Figure 6 A-D.

Leiosolenus obesus

Collections from Trincomalee, Colombo and Maggona. It can be difficult to distinguish between L. obesus and L. malaccanus. The former is often a stubby shell; the latter has a pitted concretion over the rear, a good distinguishing feature.

Continued next page

5

Maggona, Thudawa Bay, Small Boat Fishery Harbour, fishing trash.

Left - MF680: Boring in calcareous encrusted dead coral. Centre & Right - MF827:Boring into the base of a branching Tubastrea coral. Chamber de-roofed to extract shell, shown in-situ.

Leiosolenus obesus Shell flaring medially, antero-dorsal and postero-dorsal sloping; beaks terminal, inflated, with gap between beaks in dorsal view; anterior profile blunt, sub-truncate; posterior rounded, slightly narrower than anterior. Concretions thin, extension beyond posterior of shell may be present. Valentich-Scott & Tongkerd, 2008 p. 198 Figure 6 E-F

MF798: Maggona, Thudawa Bay, fishing trash, boring into coral.In situ at right.

Lithophaga Coral boring Date Mussels of the genus Lithophaga do not develop extra-periostracal concretions

MF328: Batticaloa, Kirankulam. A single valve inside the bore-hole in the dead ‘stalk’ portion of a Porites head (Image at right).

Medial part of valve showing the characteristic transverse ridges.

Lithophaga teres Chocolate date mussel Single right valve recovered from its burrow in dead coral. Inflated, chocolate brown periostracum. Hinge margin long, reaching beyond midpoint, dorsal margin not markedly angulated, anterior slightly tapered, rounded, reaching beyond beak. Fine close-set transverse ridges visible by play of light.

- Poutiers, 1998 remarks that this species bores into massive coral heads, but always below the level of live coral polyps. Also, in Abbott, 1994; Dance, 1977 p. 229;

Eisenberg, 1989 p. 156.

6

Modiolus - Horse mussels Modiolus shells have a shape described as ‘modioliform’—greatly inflated with high sail-like dorsal elevations and

bulging umbones. All are byssally attached to hard substrates.

MF186: Tangalle, Unakura Gala. Two live, heavily encrusted. Coll. Praki Thomas.

MF188: Mount Lavinia, Bellangala, byssally attached in rock crevice.

Illustrating change of shape with growth.

Modulus auriculatus Eared horse mussel Small shells with the dorsal angle in the middle or in front, the beaks level with the anterior margin and a truncate posterior. Very inflated, wider than high. Light to dark brown, hairy periostracum. Interior white or blue-white. Shape seems to change with size/age, the posterior being prolonged backwards more than height increases: compare the small triangular shell at top right with the elongated one at top left and inbetweens below it (Right-hand image). From west coast sites: Negombo, ‘Fossil Reef’, 14 m, empty on bottom; Dehiwela (Auburnside), off Godagala, empty; Wellawatte, Kinross 1st reef, 1st terrace, 3-4 m, attached Pinctada shell; Mount Lavinia, Bellangala, subtidal rocks; Maggona, Thudawa bay, SBFH, fishing trash; Tangalle, Unakura Gala. Byssally attached in rock crevices,subtidal, by diving.

MF511: Mannar, Pesalai beach, fishing trash. Bottom image of denuded shell.

MF817: Kalpitiya, Anawasala, Puttalam lagoon, fishing trash from artisanal mollusc fishery for edible meat.

Modulus modulaides Yellow banded horse muscle (FAO name) Checklist 2012 as Modiolus cf. micropterus; Modiolus metcalfei (both synonyms) Shell thin, slender, elongated or squat and trigonal. Anterior extremity in front of umbo, dorsal elevation wing-like, highest point in front of midline or behind. Posterior dorsal margin descends in a straight line to rounded posterior that is short and wide or longer and narrower. Ventral margin slightly cancave with a narrow byssal gap. Exterior covered by brown periostracum, mostly abraded over umbones and the anterior, thick and hairy over the posterior over a shell that is white ventrally, brown along the sides and purplish dorsally, or brown throughout. Interior bluish-grey or bi-color with purple upper and white lower. 63-69 mm long.

The shells variable in shape and colour—those from the sea being more slender than those from the brackish lagoon where the shells are trigonal. Both show the bi-color pattern of the interior in some of the collection. Many images of similar shell on-line.

- De Bruyne, 2007 as Modiolus micropterus; Kirtisinghe, 1978 as M. metcalfei / WoRMS MF817: Single valve, bi-color interior.

7

Modiolus philippinarum

Trincomalee, Clappenberg Bay, Sea Anglers’ Club, 1-2 m, in rock crevices, by diving. Two empty and four single valves on two

occasions, with damaged postero-dorsal and posterior margins suggestive of crab attack while in their crevices. MF183: Left and centre, one image with shell partially denuded. MF184: Right, the largest collected. Modiolus philippinarum Philippine horse mussel Large shells with a high dorsal angle behind the midline from which the postero-dorsal margin slopes down, beaks behind the anterior margin and a rounded posterior. Moderately inflated, higher than wider. White shells with a glossy brown periostracum. Interior white.

- Abbott & Dance, 1982 p. 298; Kira, 1962 p129 Pl.46#18. - As Modiolus tulipa in Kirtisinghe, 1978, a mis-identification.

Musculus cupreus

Right - MF305: Mount Lavinia, Bellangala, 7 m, under the soft coral Dendronephthea on rock. Shell denuded. Left - MF307: Mount Lavinia, Bellangala, 5 m, embedded in zooanthid tissuewith Hiatella& another species. Periostracum intact. Many specimens collected by diving at depths of 5 to 7 m. The living animals are buried in the tissues of the host near the attachment.

Musculus cupreus Thin, obliquely elliptical inflated shells with no hinge teeth. Small beaks anterior, front of shell very short, rounded, dorsal margin behind ligament extends backwards and downwards to a narrowly rounded postero-ventral angle. Ventral margin sinuate, sloping down. No byssal gape. Sculpture of growth lines crossed by two sets of weak ribs radiating from the beaks: an anterior set to the antero-ventral margin and the posterior set to the postero-ventral angle with a wide bare area inbetween: the anterior set fewer in number &arched, the posterior larger in number and narrower than the anterior. Both sets transversely ridged. Of 6 shells, 4 are coloured pale hazel with variable green tints: only the figured shell with a radiating band. Two shells much darker with chestnut concentric bands over the posterior and a darker interior. Both colour forms covered by glossy periostracum. Location data available with only one shell that was found between rock and the attaching base of a Dendronephthea. Kira says "embedded in the testa of ascidians." Most in this collection were found embedded in the bases of Dendronephthea.

- Kira, 1962 p. 127 Pl 46 #6; Bishogai Database.

8

Mytilus & Perna - Mussels These two genera are outwardly similar. They are differentiated on anatomical grounds as represented by muscle attachment scars on the shells. An anterior adductor muscle scar is present in Mytilus but not in Perna. In Mytilus the anterior and posterior components of the retractor muscle scars form a continuous band along the dorsal margin of the shell. In Perna the components attach separately (Siddal, 1980; Vakily, 1989). Three species of Perna are recognised: P. canaliculus being endemic to New Zealand, P. perna having an Atlantic distribution and P. viridis anIndo-West Pacific one (Wood et al, 2006). P. perna is also found in South India and Sri Lanka. The S. Indian community was named Perna indica(Kuriakose, 1980), later synonymised with P. perna. The muscle and retractor scars are difficult to visualize in dried shells and are of no use for the amateur collector in shell identification. See panel on page 10.

Mytilus unguiculatus Checklist 2012 as Mytilus sp. 1

Small shells with downturned beaks and straight ligament margins. Strongly angulated junction with postero-dorsal margin that is straight, slightly excavated or convex that slopes down to a broadly rounded posterior. Ventral margin straight or slightly convex. Dysodont teeth present, variable. Anterior adductor scar present. Shell horn coloured fading to white at the margins under a thin, glossy and very adherent periostracum, light brown,

MF179: Mount Lavinia, Bellangala, inshore, 3 m, attached to rock, large gregarious clumps, half buried in sand.22 shells: 51.25 - 30.36 x 27.2 - 18 mm.

with or without varying degrees of bright green behind. Interior white, pearly. As Mytilus crassitestatus Lishke in Kirtisinghe, 1978. This name and M. coruscus Gould are both synonyms.

- Lutaenko, 2005: “Widely used name Mytilus coruscus Gould, 1861, a common mussel in north-western Pacific, appeared to be a junior synonym of Mytilus unguiculatus Valenciennes, 1858.”

Left-hand image - MF180: Mount Lavinia, Bellangala rocky islet summit, intertidal, byssally attached to exposed rocks and in pools, gregarious. Other two images - MF701: Mount Lavinia, Bellangala, subtidal rocks, 2-3 m, in hollows and sides of crevices.

Continued next page

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Perna perna Common brown mussel Mytilus edulis Linnaeus in Kirtisinghe, 1978 is a mis-identification. Colour is described as “blue-black” - cleaned shells are all shades of brown.

Medium sized shells with slender, pointed anterior, the beaks pointing downward, the ligament margin weakly convex. Postero-dorsal margin straight, sloping downwards to a broadly rounded postrior. Ventral margin more or less straight or slightly concave anteriorly. Dysodont teeth small and variable in number and position. Shell light brown fading to white anteriorly and ventrally. Periostracum thin, glossy, brown, with often a green posterior margin especially on the portion reflected inside. Interior white, pearly, brownish outer colour showing through. Images show shells with eroded anteriors. Some shells have eroded dorsal angles and look sub-cylindrical. Many encrusted in life. Habitat intertidal or subtidal, usually exposed to strong wave action. Byssally attached in gregarious clumps. Collected for eating.

- Abbott & Dance, 1982 p. 297; Oliver, 1989 p. 302 (as Mytilus perna). MF376: Hambantota, Bundala, intertidal rocky shore.

Dysodont teeth of Perna perna, P. viridis and Mytilus unguiculatus can vary in number and position

In this collection the retractor scar disposition in Perna viridis is according to literature,

but not in Mytilus unguiculatus. Anterior adductor scar present in

Mytilus

10

The dysodont hinge found in the marine mussels shows a strong ligament along the

hinge line, with weak teeth near the umbones. Teeth are seen in Perna and Mytilus.

MF177: Trincomalee, Kinniya. Wild collected and cultured Clappenberg Bay for consumption at Sea Anglers’ Club.

MF816: Kalpitiya, Anawasala, Puttal-am lagoon, mollusc fishery trash.

Perna viridis Green-lipped mussel Large shells. Beaks pointing down, strongly arched ligament margin, postero-dorsal sloping down, convex, to broadly rounded posterior. Ventral margin concave. Dysodont teeth small and variable. Shells brown, fading to white at margins, anteriorly reddish, under a thick, glossy, blackish periostracum with a brilliant green margin. Shells host barnacles. Like the other mussls, after reaching a certain size, growth continues by lengthening backwards, with little or no increase in height. Research has been carried out on culture of this species in Sri Lanka. There is potential for a sustainable industry, but interest is lacking. Wild collected shells are sold.

- Kirtisinghe, 1978 p. 15; Eisenberg, 1989 p. 156; Abbott, 1994 p. 91; Oliver, 1989 p. 302; Abbott & Dance, 1982 p. 297.

Perna viridis in cultivation at Clappenberg Bay, Sea Anglers’ Club. Underwater photograph 2004.

11

Septifer Shells with strong radial sculpture and a shelly septum filling in the anterior angle behind the umbo. Byssally attached in crevices and hollows of subtidal reefs. Only one species in the country—some forms named S. virgatus in the past, now accepted as a synonym.

MF201: Mount Lavinia 1st reef, 1/2 mi south of Bellangala, 5 m, rock crevice.

A selection of shapes and colours, some deformed by growing in crevices that restrict growth.

A common species on shallow rocky reefs. Many collections from locations on the west coast: Wellawatte to Mount Lavinia, and the East coast: Trincomalee. Byssally attached in rocky crevices. Many heavily encrusted. Also found on the walls of the burrows of the sea urchin Stomopneustes virgatus on the Colombo in-shore reef. All collected by free diving.

MF194: Trincomalee, Clappenberg Bay, CSAC. 1 m, in rock crevices.

MF203: Mt Lavinia, 1st reef, south of Bellangala, 3 m, on wall of sea urchin burrow.

Septifer bilocularis Deck mussel Checklist 2012 included Septifer virgatus which is now accepted as a synonym of bilocularis. Trigonal or elongate oval. Ligament margin rises to a rounded or an angulated peak, posterior dorsal slopes down to a broadly rounded posterior. Ventral arched with hooked beaks or flattish with straight beaks. Inflated towards the ventral. Sculpture of divaricating riblets and variable growth lines. Shelf characteristic of deck mussels present behind beak. No distinctive teeth, margin crenulate. Colour variable, mostly drab orange-brown or purplish. Greens, orange or pink also seen. Interior deep blue or purplish or whitish. Periostracum thin, brownish, hairy. Shells usually heavily encrusted and maybe deformed when growing in confined spaces.

- Kirtisinghe, 1978 p. 15; Poutiers, 1998 p. 173; Eisenberg, 1989 p. 156; Abbott & Dance, 1982 p. 298.

12

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Wood, Ann R., Smita Apte, Elizabeth S. MacAvoy, Jonathan P. A. Gardner,2006. A molecular phylogeny of the marine mussel genus Perna (Bivalvia; Mytilidae) based on nuclear (ITS1 & 2) and mitochondrial (COI) DNA sequences. Molecular Phylogenetics and Evolution 44 (2007) 685-698. ScienceDirect, Elsevier. (www.elsevier.com/locate/ympev)

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Updated: 2.3.2017; 12.6.2017; 12.2.2018; 16.2.2020; 22.4.2020. 13