Hiatella Arctica Class: Bivalvia, Heterodonta, Eugeterodonta

Home , Adapedonta, Arctica (bivalve), Hiatella, Hiatella arctica, Hiatellidae, Panopea abrupta

Hiatella arctica Class: Bivalvia, Heterodonta, Eugeterodonta

Order: Imparidentia, Adapedonta A gammarid amphipod Family: Hiatelloidea, Hiatellidae

Taxonomy: There are many synonyms for Color: H. arctica due to the potentially cosmopoli- Interior: Ligament is external (Figs. 2– tan distribution of this species. Research by 3) in members of the family Hiatellidae (Coan Strauch (1968) and Beu (1969) synony- and Carlton 1975). This primary ligament mized all Hiatella species worldwide (except initially forms at the shell posterior in newly H. australis from southern Australia) as H. metamorphosed juveniles before moving to arctica (Beu 1969). Commonly seen syno- the middle of the shell (Flyachinskaya and nyms include Saxicava arctica, H. pholaids, Lezin 2008). and H. solida. Due to the extensive synony- Exterior: mizations, it is possible that there are actual- Byssus: These attachment threads ly two species currently under the name H. are present in nestling specimens, but not in arctica, locally (Coan and Valentich-Scott boring ones (e.g., H. pholadis). Adult attach 2007). with byssal threads and can also bore into rock (Coan and Valentich-Scott 2007). A Description single, long byssal thread produced by post- Size: Individuals to 50–76 mm in length larval clams allows them to be moved by (Kozloff 1993). The illustrated specimen weak water currents (see Juvenile) (Haderlie (from Coos Bay) is 38 mm in length (Quayle 1980), in a process called thread drifting (see 1970). Macoma balthica, Nutricola tantilla, Color: Exterior is white, chalky, granular, descriptions in this guide). and with tan, thin, and ragged periostracum Gills: (Hiateila, Keen and Coan 1974). The Shell: (For amino acid shell composition see interior is porcelain-like and white Brigham 1983.) Right valve slightly larger (Hiatellidae, Hunter 1949). Periostracum is than the left (Khalaman 2005) (Fig. 2). light brown or tan. Interior: Pallial line is faint and broken General Morphology: Bivalve mollusks are into discontinuous scars (Fig. 3) (Hiatellidae, bilaterally symmetrical with two lateral valves Coan and Valentich-Scott 2007), unlike or shells that are hinged dorsally and sur- Entodesma navicula (see description in this round a mantle, head, foot and viscera (see guide). Adductor muscle scars are Plate 393B, Coan and Valentich-Scott approximately equal in size, but not shape. 2007). The Veneroida is a large and diverse There is no pallial sinus (Kozloff 1974). bivalve heterodont order that is character- Exterior: The shape is highly variable ized by well-developed hinge teeth. There due to their nestling habit. Right and left val- are 22 local families, and members of the ves are equal, oblong, and gaping. The Hiatellidae are characterized by a pallial line posterior is broader and more square than the that is divided into patches (see Plate 395D, anterior end, which is broadly truncated (Fig. Coan and Valentich-Scott 2007) (Fig. 3). 1). Elongate, boring specimens have been Body: A nestling species with a thin shell reported as H. pholadis (Coan and Carlton (see Fig. 257, Kozloff 1993).

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: https://oimb.uoregon.edu/oregon-estuarine-invertebrates and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected]

Hiebert, T.C. 2015. Hiatella arctica. In: Oregon Estuarine Invertebrates: Rudys' Illustrated Guide to Common Species, 3rd ed. T.C. Hiebert, B.A. Butler and A.L. Shanks (eds.). University of Oregon Libraries and Oregon Institute of Marine Biology, Charleston, OR.

1975) (Fig. 1a). Shell sculpture is concentric abrupta and Hiatella arctica. Panopea abrup- only and the periostracum is light tan and ta tends to be larger (up to 200 mm in length) thin (Figs. 1, 2) (Hiatella, Keen and Coan than H. arctica and have a continuous pallial 1974). line, not broken into patches like in H. arctica. Hinge: Adult specimens are without Panopea abrupta, also known as the ge- (or very worn) hinge teeth (Fig. 3). Howev- oduck, is a very deep burrower with long si- er, young clams have 1–2 weak, peg-like phons (up to several feet) in soft sediments. cardinal teeth. Umbones are depressed, It has one cardinal tooth in either hinge and is nearer anterior end than middle and do not rarely found in Oregon. Also, Saxicavella touch one other (Fig. 2). pacifica, a small offshore species in soft sedi- Eyes: ments is reported (Coan and Valentich-Scott Foot: 2007). Siphons: Siphons are fused and with crim- Entodesma navicula is probably most son siphon tips (Fig. 1) (Kozloff 1993). likely to be confused with H. arctica, as it is of Burrow: a comparable size, shape, and habitat. Entodesma navicula has a dark, rough Possible Misidentifications periostracum, not a pale, thin one, an external Three bivalve families including the ligament (like H. arctica), and short, fused Lyonisiidae, Hiatellidae, and Thraciidae are siphons, but without red tips. Inside the shell characterized by their lack of dorsal margin is very pink and pearly. Entodesma navicula ears or projecting teeth or chondrophores, has no hinge teeth, but does have a large and two adductor muscles. In thraciids, the internal ligament and its pallial line is ligament can be both internal and external continuous. and the pallial line is continuous; in hiatellids Other bivalves that can be easily con- the ligament is always external and the palli- fused with H. arctica include Protothaca al line is broken into patches (see below). staminea, Petricola carditoides, Platyodon On the other hand, in the lyonisiids (e.g., cancellatus, and Cryptomya californica. The see Entodesma navicula in this guide) the venerid clam Protothaca staminea, like H. arc- pallial line is continuous, as in thraciids, but tica, is white with an external ligament, and the ligament is always internal, unlike thraci- can be found nestling in old pholad burrows. id species (Coan and Valentich-Scott 2007). It has radial as well as concentric striations, Burrowing and nestling clams, of however, and interiorly has three cardinal which there are many genera, can be diffi- hinge teeth and a strong pallial line and sinus. cult to separate by shell shape as their nestl- Petricola carditoides is a nestling clam which ing habits tend to produce a varied shell has an external ligament and a chalky white shape. Useful characteristics for differentiat- shell, as in H. arctica. It has 2–3 hinge teeth ing species include the hinge teeth, pallial in the adult, not just in the juveniles. Petricola line, and siphons. Most Pholadidae can be carditoides also has purple-tipped siphons, distinguished by their two distinct shell sec- not crimson, and its shell has some radial tions (see Penitella penita, Zirfaea pilsbryi in sculpture. The myid clam Platyodon cancella- this guide). All pholads have file-like dentic- tus is a white borer with a heavy shell with ulations and (except for Netastoma) an inter- fine, almost lamellar concentric exterior sculp- nal myophore. ture. Inside it has a chondrophore and tooth There are only two local species re- in its hinges, and a well-developed, deep pal- ported in the family Hiatellidae: Panopea lial sinus. Cryptomya californica, also a myid

clam, can nestle among rocks, although its have been collected at salinities of 30. Heart usual habitat is sand or mud. It is small (to rates 10 to 16 beats per minute at normal sa- 30 mm), thin-shelled and has a chondro- linities but drops dramatically in response to a phore. Interiorly it has an entire pallial line, change in salinity, returning to normal after 2– and an inconspicuous pallial sinus (Coan 7 days (Bakhmet et al. 2012). and Carlton 1975). Temperature: Temperature may affect shell growth and structure (see Larva) leading Ecological Information Lezin et al. (2015) to question the relevance Range: Type region is the Arcitc coast of of shell structure as taxonomic characters and Norway (Keen 1971) A circumpolar species for estimation of sea surface temperature (but with known range from Arctic Ocean to see Strauch 1968, 1971). Shell shape is also Panama (Oldroyd 1924). Range is certainly certainly affected by the nestling habits of in- influenced by many human introductions dividuals (see Rowland and Hopkins 1971). (Beu 1971; Narchi 1973; Russell-Hunter Tidal Level: Intertidal to 120 m deep, but also 1949; Yonge 197), and potentially includes low or subtidal when attached (with byssus) two species as a result (Coan and Valentich- under rocks, on floats or pilings (Morgan and Scott 2007) (see Taxonomy). In Cow Head, Allen 1976; Morton 1987; Yonge 1952, 1976; Newfoundland, radiocarbon age of H. arctica Coan and Valentich-Scott 2007). The highest fossils embedded in rock were abundance of individuals (57 per square me- approximately 8,250 years BP (Brookes and ter) was observed at 20-meter depths in the Stevens 1984). The genus Hiatella is Young Sound, northeast Greenland (Sejr et widespread in polar latitudes dating to 150 al. 2002). million years ago (Laakkonen et al. 2015). A Associates: Associates include other nestling recent molecular analysis of over 350 and boring molluscs (e.g., Entodesma navicu- specimens using three gene regions la, Penitella penita, Zirfaea pilsbryi). A poten- suggests cryptic speciation rather than tial competitor affecting settlement of Pecten widespread distribution, including at least 13 maximus, the great scallop in the Bay of Brest different putative species, several of them in France (Chauvard et al. 1996). Hiatella living sympatrically (Laakkonen et al. 2015). arctica is a facultative epibionts of the crab These authors found several distinct line- Hemigrapsus sanguineus in Japan (Isaeva et ages within the northeast Pacific (see Laak- al. 2001). Competes with and often associat- konen et al. 2015). ed with Mytilus edulis fouling communities Local Distribution: Local distribution in Co- (Khalaman 2005). Also co-occurs with the os Bay including Pigeon Point. solitary ascidian Styela rustica, in the White Habitat: Individuals nestle in old pholad bur- Sea (Khalaman 2007). rows or bore into smooth, soft, homogenous Abundance: Not common locally, however, rocks. They are also found in mussel (e.g., H. arctica is the dominant byssal bivalve in Mytilus) clumps, on pilings, and on open the arctic and boreal regions (Coan and coasts within algal holdfasts. On hard Valentich-Scott 2007). Along with Mytilus surfaces and within crevices, individuals edulis, H. arctica was the most abundant bi- attach byssally (Hunter 1949). Prefers valves in Eyjafordur, North Iceland, with newly sheltered locations or being covered by metamorphosed spat observed on settlement other organisms (Khalaman 2005). plates throughout the year (Garcia et al. Salinity: Occurs in Coos Bay as well as 2008). Density reached approximately 815 more saline parts of estuary, and individuals individuals per meter in the White Sea

(Khalaman 2005). margins. Veliger larvae of H. arctica have a distinctly triangular shell and there are two Life-History Information distinct posterior dorsal spines on the disso- Reproduction: Spawning occurs from mid conch in newly metamorphosed H. arctica June through September and veliger larvae (see Fig. 3F, Flyachinskaya and Lezin 2006). are present through November (White Sea, These spines are lost in adulthood Flya- Russia, Flyachinskaya and Lesin 2006). chinskaya and Lezin 2008). Settlement was The early development is apparently very observed throughout the year in Eyjafjordur, similar to Mytilus edulis (see description in Iceland with peaks in abundance of primary this guide), which was described by Mala- (<1mm) settlers in September. Lezin et al. khov and Medvedeva (1985). (2015) found that larvae and juveniles raised Larva: Bivalve development generally pro- at 12˚C formed ridges and spines on the shell ceeds from external fertilization via broad- posterior. However, they developed fewer or cast spawning through a ciliated trocho- inconspicous spines at lower temperatures phore stage to a veliger larva. Bivalve veli- (e.g., 5˚C) (Lezin et al. 2015). Wild-caught gers are characterized by a ciliated velum planktonic larvae measured up to 380 µm in that is used for swimming, feeding and respi- the North Sea and had a tough shell, a con- ration. The veliger larva is also found in spicuous pallial line, and concentric ridges many gastropod larvae, but the larvae in the that become more pronounced and widely two groups can be recognized by shell mor- spaced with distance from the umbo (Rees phology (i.e. snail-like versus clam-like). In 1950). bivalves, the initial shelled-larva is called a D Juvenile: Like other bivalve species (e.g. -stage or straight-hinge veliger due to the Mytilus edulis, Macoma balthica), post-larval “D” shaped shell. This initial shell is called a (i.e., juvenile) H. arctica may exploit two dis- prodissoconch I and is followed by a prodis- persal periods. The initial larval dispersal and soconch II, or shell that is subsequently add- settlement into the benthos, is followed by po- ed to the initial shell zone (see Fig. 1, Caddy tential dispersal from the benthos by thread 1969). Finally, shell secreted following met- drifting on water currents via mucous threads amorphosis is simply referred to as the dis- by juveniles (i.e., “thread drifting”, Martel and soconch (see Fig. 2, Brink 2001). Once the Chia 1991). larva develops a foot, usually just before Longevity: Age was estimated for individuals metamorphosis and loss of the velum, it is in the Young Sound, northeast Greenland by called a pediveliger (see Fig. 1, Caddy 1969; counting growth rings (a method validated by Kabat and O’Foighil 1987; Brink 2001). (For Sejr et al. 200b); the oldest individuals were generalized life cycle see Fig. 1, Brink estimated to be 126 years old (Sejr et al. 2001). Larval development was described 2002a). by Flyachinskaya and Lesin (2006) where Growth Rate: Mean growth rate was estimat- straight hinge larvae were 120 µm, umbo ed to be 0.14 mm per year in northeast were 160 µm, early pediveligers were 220 Greenland (Sejr et al. 2002a). µm, pediveligers were 310 µm and grew to Food: A suspension feeder. Filtration rates 400 µm pre-metamorphosis and were 750 were 1.4 x 10-2 liters per hour per gram wet µm post-metamorphosis (see Fig. 3, Flya- weight (Phaeodactylum tricornutum, 15˚C) chinskaya and Lesin 2006). The shell is yel- and increases up to temperatures of 17˚C be- lowish pink when early in development and fore decreasing above 25˚C (Ali 1970), at high changes to a darker (dirtier) yellow with pink temperatures H. arctica close their valves

completely (Petersen et al. 2003). Clear- 129-149. In: Identification guide to larval ance rates for individuals at varying temper- marine invertebrates of the Pacific North- atures were recorded by Petersen et al. west. A. Shanks (ed.). Oregon State Uni- (2003) for individuals collected in the Arctic versity Press, Corvallis, OR. (Young Sound, northeast Greenland) and 6. BROOKES, I. A., and R. K. STEVENS. temperate waters (Tjarno, Sweden) (see Ta- 1985. Radiocarbon age of rock boring Hia- ble 1, Petersen et al. 2003) and averages tella arctica (Linne) and postglacial sea being approximately 23.4 and 27.4 ml per level change at Cow-Head, Newfoundland. minute per gram body weight those from the Canadian Journal of Earth Sciences. Arctic and Sweden, respectively (Petersen 22:136-140. et al. 2003). Clearance rates and assimila- 7. CHAUVAUD, L., G. THOUZEAU, and J. tion efficiency (how much food is metabo- GRALL. 1996. Experimental collection of lized) of Rhodomonas baltica decreases great scallop postlarvae and other benthic with increasing food concentration (Sejr et species in the Bay of Brest: Settlement al. 2004). Sejr et al. (2004) found that indi- patterns in relation to spatio-temporal vari- viduals grow to annual observed growth in ability of environmental factors. Aquacul- the wild in three weeks at optimal feeding ture International. 4:263-288. conditions in the laboratory and suggest that 8. COAN, E. V., and J. T. CARLTON. 1975. food is limiting in wild populations. Phylum Mollusca: Bivalvia, p. 543-578. In: Predators: Toothed snails (e.g., Nucella Light's manual; intertidal invertebrates of spp.) can prey on small nestling clams like the central California coast. S. F. Light, R. H. arctica. I. Smith, and J. T. Carlton (eds.). Universi- Behavior: Individuals bore mechanically ty of California Press, Berkeley. and without the aid of chemical compounds 9. COAN, E. V., and P. VALENTICH-SCOTT. (Hunter 1949). 2007. Bivalvia, p. 807-859. In: The Light and Smith manual: intertidal invertebrates Bibliography from central California to Oregon. J. T. 1. ALI, R. M. 1970. Influence of suspension Carlton (ed.). University of California density and temperature on filtration rate Press, Berkeley, CA. of Hiatella arctica. Marine Biology. 6:291- 10. FLYACHINSKAYA, L. P., and P. A. LE- 302. SIN. 2006. Using 3D reconstruction meth- 2. BAKHMET, I. N., A. J. KOMENDANTOV, od in the investigations of bivalvia larval and A. O. SMUROV. 2012. Effect of sa- development (by the example of Hiatella linity change on cardiac activity in Hiatel- arctica L.). Trudy Zoologicheskogo Insti- la arctica and Modiolus modiolus, in the tuta. 310:45-50. White Sea. Polar Biology. 35:143-148. 11. FLYACHINSKAYA, L. P., and P. A. LEZIN. 3. BEU, A. G. 1971. New light on variation 2008. Larval and juvenile shell develop- and taxonomy of bivalve Hiatella. New ment in the White Sea bivalve Hiatella arc- Zealand Journal of Geology and Geo- tica (Linnaeus, 1767). Zoologiya Be- physics. 14:64-66. spozvonochnykh. 5:39-46. 4. BRIGHAM, J. K. 1983. Intrashell varia- 12. GARCIA, E. G., G. G. THORARINSDOT- tions in amino acid concentrations and TIR, and S. A. RAGNARSSON. 2003. Set- isoleucine epimerization ratios in fossil tlement of bivalve spat on artificial collec- Hiatella arctica. Geology. 11:509-513. tors in Eyjafjordur, North Iceland. Hydrobi- 5. BRINK, L. A. 2001. Mollusca: Bivalvia, p. ologia. 503:131-141.

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