Macoma Nasuta Class: Bivalvia; Heterodonta Order: Veneroida the Bent Nosed Clam Family: Tellinidae

Phylum: Mollusca Macoma nasuta Class: Bivalvia; Heterodonta Order: Veneroida The bent nosed clam Family: Tellinidae

Taxonomy: Originally described in the genus you, the right valve is in the right hand (Fig. 4) Tellina by Conrad in 1837, additional (Keen and Coan 1974). synonyms include Tellina tersa, Macoma Body: kelseyi, M. jacalitosana. Also subspecific Color: designations (e.g. Macoma Heteromacoma Interior: Ligament is entirely external nasuta, Kabat and O’Foighil 1987) are and the is not supported by a nymph or sometimes seen. However, M. nasuta is the projection dorsally (Fig. 5). name almost exclusively used in current Exterior: intertidal guides (e.g. Coan and Valentich- Byssus: Scott 2007). Gills: Shell: Shells ovate (Fig. 1) and posterior Description portions of valves are distinctly bent to the Size: Individuals range from 3 to 70 mm right (“bent nose”, Kozloff 1993) (Fig. 4). The (Macginitie and Macginitie 1949; Kozloff anterior end is rounded, and the posterior is 1993) and are seldom larger than 64 mm wedge-shaped, or truncate (i.e. not flanged). (Packard 1918). In Coos Bay, the largest Interior: The pallial sinus of the right individuals are approximately 58 mm, which valves does not reach the anterior adductor would classify them as having a medium scar (Fig. 3) (see Plate 422, Coan and sized shell (see Keen and Coan 1974). Valentich-Scott 2007). The adductor and Color: Shell is white and chalky where posterior muscle scars are similar in shape in eroded (see Fig. 293, Kozloff 1993) and has both valves and overlaps, but sinus patterns dark brown parchment periostracum, differ. The pallial sinus of the left valve especially near lower edge and siphons on reaches the anterior adductor muscle scar, valves. Shell is often with black markings fuses and overlaps with it (Fig. 2) (Coan and externally (Brusca and Brusca 1978) but there Valentich-Scott 2007). is no interior shell color (Keen and Coan Exterior: Valve exterior is thin and 1974). Siphons can be orange (Kozloff smooth, but not polished. The shells are 1993). sometimes blackish and are thin, with fine General Morphology: Bivalve mollusks are radial lines. bilaterally symmetrical with two lateral valves Hinge: Hinge with ligament is entirely or shells that are hinged dorsally and external and no lateral teeth (Fig. 5) surround a mantle, head, foot and viscera (Macoma, Coan and Valentich-Scott 2007). (see Plate 393B, Coan and Valentich-Scott Two cardinal hinge teeth are present on the 2007). Among the bivalves, the Heterodonta right valve (Fig. 5) and one on the left valve are characterized by ctenidia (or gills) that (Fig. 2). The beak is central and slightly are eulamellibranchiate, fused mantle prominent (Fig. 5) (Packard 1918). margins and the presence of long siphons. Eyes: Veneroid bivalves have well-developed hinge Foot: teeth and members of the family Tellinidae Siphons: The inhalant and exhalant siphons have short lateral hinge teeth (when present – are completely separate and distinctly orange see Possible Misidentifications), shells with in color (Fig. 6) (Kozloff 1993). external striations or ribs, and deep pallial Burrow: Clams burrow 10–20 cm within fine sinuses (Coan and Valentich-Scott 2007). sediment, but these burrows are not When holding closed shell in both hands with permanent (Alexander et al. 1993; the hinged area up and the ligaments toward Volkenborn et al. 2012). For burrow

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12909 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] schematic, see Fig. 8.10B, Zonneveld et al. extending from posterior end to the external 2014. ligament, this is absent in other Macoma species. The former species is called the the Possible Misidentifications sand clam and has a quadrate and truncate Tellinids can be distinguished from posterior. The latter is elongate, has a other small or young bay clams (i.e. pointed posterior, unique muscle scars, is Mactridae: Tresus; Veneridae: Protothaca, relatively rare and small (to 2.5 cm) and Saxidomus; Myidae: Mya, Cryptomya) an occurs from Trinidad, California southward. external ligament, an ovate shell, an Macoma secta, also has a white shell, with a inconspicuous nymph (or supporting yellowish epidermis. Its right valve is more projection for the external ligament), inflated than the left, and it can be large (to sometimes reddish hue and lateral teeth as 120 mm) and is more common in clean sand, well as a shell with ribs or striations (no radial not in estuarine mud. pattern) and shells that never gape. Lateral The morphology of the pallial sinus teeth may or may not be present in the differentiates the other species. In species Tellinidae (Coan 1971). Myids have a hinge without a posterior dorsal flange, M. acolasta with a spoon-shaped chondrophore (left and M. yoldiformis, the anterior ventral edge valve) and a projecting tooth (right valve) (see of the pallial sinus is detached for a portion of Mya arenaria, this guide). Venerids have the distance to the posterior adductor muscle three cardinal teeth in each valve. Mactrids scar. Macoma acolasta also has a rounded have an internal ligament, A-shaped cardinal posterior, rather than pointed as in M. teeth, and gaping valves (Coan and yoldiformis and is rare, sand-dwelling, and Valentich-Scott 2007). The Tellinidae has occurs from Bodega Bay, California, around 16 species distributed between two southward. Macoma yoldiformis is elongate, genera locally – Tellina and Macoma. These inflated, and thin, with the pallial sinus genera can be differentiated by the hinge detached from the pallial line. Although the teeth, Tellina species have a hinge with range of this clam is from Vancouver south to lateral teeth, while Macoma species do not. Baja California, it is not included in Puget Macoma species have shells that are also Sound or British Columbia work (Dunnill and more rounded and inflated thanTellina, and Ellis 1969). It can be found in silt in low are smooth, white, often chalky. They are intertidal of protected bays (Coan and characterized by having a ovate shell Valentich-Scott 2007). consisting of two equal valves, a dark and Macoma inquinata, M. nasuta and M. deciduous periostracum, two cardinal teeth, balthica (see descriptions in this guide) are all the absence of lateral teeth and a pallial sinus species with an anterior ventral edge of pallial that is deeper on the left valve (Scott and sinus that is not detached and they tend to be Blake 1998; Arruda and Domaneschi 2005). larger (up to 110 mm) than M. acolasta or M. Macoma species may also have a more yoldiformis (less than 30 mm). Macoma northern geographic distribution whileTellina balthica has a pinkish hue and a pallial sinus are elongate, relatively compressed, that ends ¾ of the way to anterior adductor conspicuously sculptured, brightly colored, muscle scar and is generally more oval than and usually warm water dwellers (Coan M. nasuta or M. inquinata (Kozloff 1993). 1971). Eleven species in the infaunal genus Macoma inquinata is a common mud clam, Macoma (Luttikhuizen et al. 2012) are with slightly inflated but not dramatically bent reported locally (although 30 have been valves. In M. inquinata, the pallial sinus does identified in the northeastern Pacific, Dunnill not reach the ventral end of the anterior and Ellis 1969), but only seven are described adductor muscle and the shell is chalky white in local keys (e.g. Coan and Valentich-Scott with a fibrous olive green periostracum. 2007), the four most common species of Macoma nasuta, on the other hand, is not as Macoma in our area are M. balthica, M. round and heavy as M. inquinata and its nasuta, M. inquinata, and M. secta (Kabat pallial sinus reaches and joins the anterior and O’Foighil 1987). adductor scar above its base (left valve). (Its Two species, M. secta and M. right valve may be more like M. inquinata’s). indentata have a posterior dorsal flange Furthermore, its siphons are orange and its Hiebert, T.C. 2015. Macoma nasuta. 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. shell posterior is bent to the right (Fig. 4). muddy and silty sediment (i.e., a wide range Macoma inquinata can also bend slightly of grain sizes, Alexander et al. 1993) up to 40 posteriorly, and may be confused with the cm (Rae 1978). Macoma nasuta was the thinner M. nasuta, without investigations of oldest intertidal species present at Queen the other aforementioned features. In M. Charlotte Islands in the fossil record (~13,210 balthica, the pallial sinus that reaches to 1/4 C14 years) and was possibly the first species the anterior adductor muscle scar and the to colonize this area following the last glacial shell has a pinkish hue. (see Plate 422 for maximum. This early colonization was likely diagrams of these distinguishing due to the tolerance of M. nasuta for low characteristics in Macoma). Macoma balthica water temperature, high turbidity, low primary and M. inquinata are generally smaller than productivity and low salinity. Following initial M. nasuta (up to 5 cm) and the shell of M. colonization, congeneric species, M. calcarea nasuta is white in-side and out, with some and M. inquinata, began to appear in the dark periostracum. fossil record (Hetherington and Reid 2003). (The following species may be present Salinity: Adapted to a wide range of locally, but are not included in local conditions and salinities. dichotomous keys). Macoma expansa, is a Temperature: Temperate and cold waters. rare, usually offshore species (to 50 mm) Macoma nasuta is not found in the Panamic whose pallial sinuses are perpendicular to the province to the south. pallial line. Macoma elimata is found only in Tidal Level: Intertidal and subtidal (Kabat 15–476 meters of water. Macoma incongrua and O’Foighil 1987). Most common in bays at is a generally northern species which can be mid-tide line (Coan and Valentich-Scott found to 33°N latitude, intertidally to 36 2007). Also reported from low tide horizon to meters. It has somewhat inflated valves, is 137 meters, with decreasing numbers with usually 30–40 mm long, and almost round in depth (Rae 1978). outline (Dunnill and Ellis 1969). Macoma Associates: Occasionally infested with calcarea is found from 35 meters and lower, encysted larvae of the tapeworm from 37° north. Other northern subtidal Anthobothrium sp. (MacGinitie and MacGinitie species include the large M. brota and M. 1949). Juvenile pea crabs, Pinnixia littoralis, lipara (Dunnill and Ellis 1969). can occur within the mantle cavity of M. inquinata and M. nasuta in Puget Sound, Ecological Information Washington (Haderlie and Abbott 1980). Also Range: Type locality is Astoria, OR. Known host to the commensal nemertean range from Kodiak, Alaska to Baja California Malacobdella macomae (Haderlie and Abbott (Ricketts and Calvin 1971). Kodiak Island, 1980; Kozloff 1991; Roe et al. 2007). Alaska to Cabo San Lucas, Baja California Individuals also host the turbellarian, Graffilla (Coan 1971; Rae 1978). pugetensis in the pericardial cavity (Schell Local Distribution: Occurs locally, in bays 1989). While other bivalve species exhibited as well as offshore below the surf zone (Coan declined numbers associated with the exotic and Valentich-Scott 2007). mussel, Musculista senhousia, M. nasuta Habitat: Preferred substrates include mud populations were not negatively affected and, and muddy sand (Kabat and O’Foighil 1987), instead, increased in number (Mission Bay, about 10–15 cm below the surface (Kozloff San Diego, California, Crooks 2001). 1993). Macoma nasuta is very adaptable and Macoma nasuta co-occurs with the congener, can live in finer mud than other Macoma M. inquinata. species, often in the extremely stale waters of Abundance: On “every possible mud flat" small lagoons (Ricketts and Calvin 1971). (Ricketts and Calvin 1971) where it is often Individuals also found in eelgrass beds the most common clam (e.g., in Elkhorn (Kozloff 1974). A common deposit feeder is Slough). Individuals are more abundant in often used in sediment toxicity testing and habitats with more mud than sand (San biomarker responses (e.g., Boese et al. 1995; Francisco Bay, California, Alexander et al. Werner et al. 2004; Cho et al. 2007; 1993). The most abundant bivalve collected Amirbahman et al. 2013). Can burrow within in box cores (25 x 25 x 50 cm) from a coastal

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12909 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] lagoon in Bodega Harbor, California (Everett Food: Primarily a suspension feeder that 1994). also ingests detritus from mud surface with siphon, while discarding coarse, inedible Life-History Information material (MacGinitie and MacGinitie 1949; Reproduction: Separate sexes, gametes are Kabat and O’Foighil 1987). In a comparison discharged into the water through excurrent of individual growth on a diet of surface siphon. Oregon spawning reportedly spring, sediment versus detritus, the former early summer (Haderlie and Abbott 1980). supported better growth (Hylleberg and Nuclear protein composition of sperm is Gallucci 1975). When deposit feeding, M. described by Ausio (1988). Gametogenesis nasuta sucks the top 1 mm of sediment by for M. nasuta and M. secta is described by boring or rotating with the siphon tip (Fig. 6) Rae (1978) and reproduction and and exhibits some selectivity in feeding development has been described for the (Hylleberg and Gallucci 1975; Gallucci and common congener, M. balthica (Caddy 1967, Hylleberg 1976). Interstitial water amounted 1969; Lammens 1967). Unspawned oocytes to only 4% of the total water ventilated (~0.2 of M. nasuta were 48–57 µm in diameter. ml per hour). Instead, the overlying water is Individuals were observed to spawn most primarily ventilated (~7.3 ml per hour, Winsor months of the year, with peak spawning from 1990). The average ventilation rate for M. August to November (14˚C, Rae 1978). nasuta was found to be lower than that of Spawning occurs in a a brief period of two other filtering bivalves, suggesting that days and sex ratios were 1:1, with no deposit feeding may be their dominant hermaphrodites seen in the populations feeding mode (Specht and Lee 1989). (Tamales Bay, California, Rae 1978). Macoma nasuta was found to be actively Larva: Bivalve development generally deposit feeding 83% of the time (Volkenborng proceeds from external fertilization via et al. 2012). (For irrigation patterns see Table broadcast spawning through a ciliate 1, Volkenborn et al. 2012). The deposit trochophore stage to a veliger larva. Bivalve feeding behavior of both M. nasuta and M. veligers are characterized by a ciliated velum inquinata showed variation in response to that is used for swimming, feeding and water flow. Their inhalant siphons extended respiration. The veliger larva is also found in farther, allowing for deposit feeding over a many gastropod larvae, but the larvae in the larger area, when water flow was lower two groups can be recognized by shell (Levinton 1991). morphology (i.e. snail-like versus clam-like). Predators: Small clams are fed upon by In bivalves, the initial shelled-larva is called a crabs (e.g., Cancer productus), seastars (e.g., D-stage or straight-hinge veliger due to the Pisaster spp.), as well as the snail Polinices “D” shaped shell. This initial shell is called a lewisii (Haderlie and Abbott 1980; Kozloff prodissoconch I and is followed by a 1993) and shore birds. prodissoconch II, or shell that is subsequently Behavior: Usually situated within the mud added to the initial shell zone (e.g. see M. with left valve down (MacGinitie and balthica, Fig. 1, Caddy 1969). Finally, shell MacGinitie 1949; Kozloff 1993) and burrows secreted following metamorphosis is simply 10–20 cm deep into the sediment referred to as the dissoconch (see Fig. 2, (Volkenborn et al. 2012) with a burrowing rate Brink 2001). Once the larva develops a foot, that is highest in fine sand and muddy silt usually just before metamorphosis and loss of (Alexander et al. 1993). the velum, it is called a pediveliger (Kabat and O’Foighil 1987; Brink 2001). (For generalized Bibliography life cycle see Fig. 1, Brink 2001). Macoma nasuta and M. secta are known to have free 1. ALEXANDER, R. R., R. J. STANTON, swimming veliger larvae (MacGinitie and and J. R. DODD. 1993. Influence of MacGinitie 1949; Marriage 1954; Rae 1978, sediment grain-size on the burrowing 1979; Brink 2001). of bivalves: correlation with distribution Juvenile: and stratigraphic persistence of Longevity: selected neogene clams. Palaios. Growth Rate: 8:289-303. Hiebert, T.C. 2015. Macoma nasuta. 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