Macoma Nasuta Class: Bivalvia, Heterodonta, Euheterodonta

Macoma nasuta Class: Bivalvia, Heterodonta, Euheterodonta

Order: Imparidentia, Cardiida The bent nosed clam Family: Tellinoidea, Tellinidae, Macominae

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

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. 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.

right valve (Fig. 5) and one on the left valve teeth and a pallial sinus that is deeper on the (Fig. 2). The beak is central and slightly left valve (Scott and Blake 1998; Arruda and prominent (Fig. 5) (Packard 1918). Domaneschi 2005). Macoma species may Eyes: also have a more northern geographic distri- Foot: bution whileTellina are elongate, relatively Siphons: The inhalant and exhalant siphons compressed, conspicuously sculptured, are completely separate and distinctly brightly colored, and usually warm water dwe- orange in color (Fig. 6) (Kozloff 1993). llers (Coan 1971). Eleven species in the in- Burrow: Clams burrow 10–20 cm within fine faunal genus Macoma (Luttikhuizen et al. sediment, but these burrows are not perma- 2012) are reported locally (although 30 have nent (Alexander et al. 1993; Volkenborn et been identified in the northeastern Pacific, al. 2012). For burrow schematic, see Fig. Dunnill and Ellis 1969), but only seven are 8.10B, Zonneveld et al. 2014. described in local keys (e.g. Coan and Valen- tich-Scott 2007), the four most common spe- Possible Misidentifications cies of Macoma in our area are M. balthica, Tellinids can be distinguished from M. nasuta, M. inquinata, and M. secta (Kabat other small or young bay clams (i.e. and O’Foighil 1987). Mactridae: Tresus; Veneridae: Protothaca, Two species, M. secta and M. inden- Saxidomus; Myidae: Mya, Cryptomya) an tata have a posterior dorsal flange extending external ligament, an ovate shell, an incons- from posterior end to the external ligament, picuous nymph (or supporting projection for this is absent in other Macoma species. The the external ligament), sometimes reddish former species is called the the sand clam hue and lateral teeth as well as a shell with and has a quadrate and truncate posterior. ribs or striations (no radial pattern) and she- The latter is elongate, has a pointed posterior, lls that never gape. Lateral teeth may or unique muscle scars, is relatively rare and may not be present in the Tellinidae (Coan small (to 2.5 cm) and occurs from Trinidad, 1971). Myids have a hinge with a spoon- California southward. Macoma secta, also shaped chondrophore (left valve) and a pro- has a white shell, with a yellowish epidermis. jecting tooth (right valve) (see Mya arenaria, Its right valve is more inflated than the left, this guide). Venerids have three cardinal and it can be large (to 120 mm) and is more teeth in each valve. Mactrids have an inter- common in clean sand, not in estuarine mud. nal ligament, A-shaped cardinal teeth, and The morphology of the pallial sinus dif- gaping valves (Coan and Valentich-Scott ferentiates the other species. In species with- 2007). The Tellinidae has around 16 spe- out a posterior dorsal flange, M. acolasta and cies distributed between two genera locally M. yoldiformis, the anterior ventral edge of the – Tellina and Macoma. These genera can pallial sinus is detached for a portion of the be differentiated by the hinge teeth, Tellina distance to the posterior adductor muscle species have a hinge with lateral teeth, while scar. Macoma acolasta also has a rounded Macoma species do not. Macoma species posterior, rather than pointed as in M. yoldi- have shells that are also more rounded and formis and is rare, sand-dwelling, and occurs inflated thanTellina, and are smooth, white, from Bodega Bay, California, southward. often chalky. They are characterized by ha- Macoma yoldiformis is elongate, inflated, and ving a ovate shell consisting of two equal thin, with the pallial sinus detached from the valves, a dark and deciduous periostracum, pallial line. Although the range of this clam is two cardinal teeth, the absence of lateral from Vancouver south to Baja California, it is

not included in Puget Sound or British rare, usually offshore species (to 50 mm) Columbia work (Dunnill and Ellis 1969). It whose pallial sinuses are perpendicular to can be found in silt in low intertidal of the pallial line. Macoma elimata is found protected bays (Coan and Valentich-Scott only in 15–476 meters of water. Macoma 2007). incongrua is a generally northern species Macoma inquinata, M. nasuta and M. which can be found to 33°N latitude, balthica (see descriptions in this guide) are intertidally to 36 meters. It has somewhat all species with an anterior ventral edge of inflated valves, is usually 30–40 mm long, pallial sinus that is not detached and they and almost round in outline (Dunnill and Ellis tend to be larger (up to 110 mm) than M. ac- 1969). Macoma calcarea is found from 35 olasta or M. yoldiformis (less than 30 mm). meters and lower, from 37° north. Other Macoma balthica has a pinkish hue and a northern subtidal species include the large pallial sinus that ends ¾ of the way to anteri- M. brota and M. lipara (Dunnill and Ellis or adductor muscle scar and is generally 1969). more oval than M. nasuta or M. inquinata Ecological Information (Kozloff 1993). Macoma inquinata is a Range: Type locality is Astoria, OR. Known common mud clam, with slightly inflated but range from Kodiak, Alaska to Baja California not dramatically bent valves. In M. (Ricketts and Calvin 1971). Kodiak Island, inquinata, the pallial sinus does not reach Alaska to Cabo San Lucas, Baja California the ventral end of the anterior adductor (Coan 1971; Rae 1978). muscle and the shell is chalky white with a Local Distribution: Occurs locally, in bays as fibrous olive green periostracum. Macoma well as offshore below the surf zone (Coan nasuta, on the other hand, is not as round and Valentich-Scott 2007). and heavy as M. inquinata and its pallial Habitat: Preferred substrates include mud sinus reaches and joins the anterior and muddy sand (Kabat and O’Foighil 1987), adductor scar above its base (left valve). (Its about 10–15 cm below the surface (Kozloff right valve may be more like M. inquinata’s). 1993). Macoma nasuta is very adaptable and Furthermore, its siphons are orange and its can live in finer mud than other Macoma shell posterior is bent to the right (Fig. 4). species, often in the extremely stale waters of Macoma inquinata can also bend slightly small lagoons (Ricketts and Calvin 1971). In- posteriorly, and may be confused with the dividuals also found in eelgrass beds (Kozloff thinner M. nasuta, without investigations of 1974). A common deposit feeder is often the other aforementioned features. In M. used in sediment toxicity testing and balthica, the pallial sinus that reaches to 1/4 biomarker responses (e.g., Boese et al. 1995; the anterior adductor muscle scar and the Werner et al. 2004; Cho et al. 2007; shell has a pinkish hue. (see Plate 422 for Amirbahman et al. 2013). Can burrow within diagrams of these distinguishing muddy and silty sediment (i.e., a wide range characteristics in Macoma). Macoma of grain sizes, Alexander et al. 1993) up to 40 balthica and M. inquinata are generally cm (Rae 1978). Macoma nasuta was the smaller than M. nasuta (up to 5 cm) and the oldest intertidal species present at Queen shell of M. nasuta is white in-side and out, Charlotte Islands in the fossil record (~13,210 with some dark periostracum. C14 years) and was possibly the first species (The following species may be pre- to colonize this area following the last glacial sent locally, but are not included in local maximum. This early colonization was likely dichotomous keys). Macoma expansa, is a

due to the tolerance of M. nasuta for low lected in box cores (25 x 25 x 50 cm) from a water temperature, high turbidity, low coastal lagoon in Bodega Harbor, California primary productivity and low salinity. (Everett 1994). Following initial colonization, congeneric Life-History Information species, M. calcarea and M. inquinata, Reproduction: Separate sexes, gametes are began to appear in the fossil record discharged into the water through excurrent (Hetherington and Reid 2003). siphon. Oregon spawning reportedly spring, Salinity: Adapted to a wide range of condi- early summer (Haderlie and Abbott 1980). tions and salinities. Nuclear protein composition of sperm is de- Temperature: Temperate and cold waters. scribed by Ausio (1988). Gametogenesis for Macoma nasuta is not found in the Panamic M. nasuta and M. secta is described by Rae province to the south. (1978) and reproduction and development Tidal Level: Intertidal and subtidal (Kabat has been described for the common conge- and O’Foighil 1987). Most common in bays ner, M. balthica (Caddy 1967, 1969; Lam- at mid-tide line (Coan and Valentich-Scott mens 1967). Unspawned oocytes of M. nasu- 2007). Also reported from low tide horizon ta were 48–57 µm in diameter. Individuals to 137 meters, with decreasing numbers were observed to spawn most months of the with depth (Rae 1978). year, with peak spawning from August to No- Associates: Occasionally infested with en- vember (14˚C, Rae 1978). Spawning occurs cysted larvae of the tapeworm Anthobothri- in a a brief period of two days and sex ratios um sp. (MacGinitie and MacGinitie 1949). were 1:1, with no hermaphrodites seen in the Juvenile pea crabs, Pinnixia littoralis, can populations (Tamales Bay, California, Rae occur within the mantle cavity of M. 1978). inquinata and M. nasuta in Puget Sound, Larva: Bivalve development generally pro- Washington (Haderlie and Abbott 1980). ceeds from external fertilization via broadcast Also host to the commensal nemertean spawning through a ciliate trochophore stage Malacobdella macomae (Haderlie and to a veliger larva. Bivalve veligers are charac- Abbott 1980; Kozloff 1991; Roe et al. 2007). terized by a ciliated velum that is used for Individuals also host the turbellarian, swimming, feeding and respiration. The veli- Graffilla pugetensis in the pericardial cavity ger larva is also found in many gastropod lar- (Schell 1989). While other bivalve species vae, but the larvae in the two groups can be exhibited declined numbers associated with recognized by shell morphology (i.e. snail-like the exotic mussel, Musculista senhousia, M. versus clam-like). In bivalves, the initial nasuta populations were not negatively shelled-larva is called a D-stage or straight- affected and, instead, increased in number hinge veliger due to the “D” shaped shell. (Mission Bay, San Diego, California, Crooks This initial shell is called a prodissoconch I 2001). Macoma nasuta co-occurs with the and is followed by a prodissoconch II, or shell congener, M. inquinata. that is subsequently added to the initial shell Abundance: On “every possible mud zone (e.g. see M. balthica, Fig. 1, Caddy flat" (Ricketts and Calvin 1971) where it is 1969). Finally, shell secreted following meta- often the most common clam (e.g., in morphosis is simply referred to as the disso- Elkhorn Slough). Individuals are more abun- conch (see Fig. 2, Brink 2001). Once the lar- dant in habitats with more mud than sand va develops a foot, usually just before meta- (San Francisco Bay, California, Alexander et morphosis and loss of the velum, it is called a al. 1993). The most abundant bivalve col-

pediveliger (Kabat and O’Foighil 1987; Brink Kozloff 1993) and shore birds. 2001). (For generalized life cycle see Fig. 1, Behavior: Usually situated within the mud Brink 2001). Macoma nasuta and M. secta with left valve down (MacGinitie and are known to have free swimming veliger MacGinitie 1949; Kozloff 1993) and burrows larvae (MacGinitie and MacGinitie 1949; 10–20 cm deep into the sediment (Volkenborn Marriage 1954; Rae 1978, 1979; Brink et al. 2012) with a burrowing rate that is 2001). highest in fine sand and muddy silt (Alexander Juvenile: et al. 1993). Longevity: Bibliography Growth Rate: Food: Primarily a suspension feeder that 1. ALEXANDER, R. R., R. J. STANTON, also ingests detritus from mud surface with and J. R. DODD. 1993. Influence of sedi- siphon, while discarding coarse, inedible ment grain-size on the burrowing of bi- material (MacGinitie and MacGinitie 1949; valves: correlation with distribution and Kabat and O’Foighil 1987). In a comparison stratigraphic persistence of selected neo- of individual growth on a diet of surface sedi- gene clams. Palaios. 8:289-303. ment versus detritus, the former supported 2. AMIRBAHMAN, A., D. I. MASSEY, G. LO- better growth (Hylleberg and Gallucci 1975). TUFO, N. STEENHAUT, L. E. BROWN, J. When deposit feeding, M. nasuta sucks the M. BIEDENBACH, and V. S. MAGAR. top 1 mm of sediment by boring or rotating 2013. Assessment of mercury bioavailabil- with the siphon tip (Fig. 6) and exhibits some ity to benthic macroinvertebrates using dif- selectivity in feeding (Hylleberg and Gallucci fusive gradients in thin films (DGT). Envi- 1975; Gallucci and Hylleberg 1976). Intersti- ronmental Science-Processes & Impacts. tial water amounted to only 4% of the total 15:2104-2114. water ventilated (~0.2 ml per hour). Instead, 3. ARRUDA, E. P., and O. DOMANESCHI. the overlying water is primarily ventilated 2005. New species of Macoma (Bivalvia: (~7.3 ml per hour, Winsor 1990). The aver- Tellinoidea: Tellinidae) from southeastern age ventilation rate for M. nasuta was found Brazil, and with description of its gross to be lower than that of other filtering bi- anatomy. Zootaxa:13-22. valves, suggesting that deposit feeding may 4. AUSIO, J. 1988. An unusual cysteine- be their dominant feeding mode (Specht and containing histone H1-like protein and 2 Lee 1989). Macoma nasuta was found to be protamine-like proteins are the major nu- actively deposit feeding 83% of the time clear proteins of the sperm of the bivalve (Volkenborng et al. 2012). (For irrigation mollusk Macoma nasuta. Journal of Bio- patterns see Table 1, Volkenborn et al. logical Chemistry. 263:10141-10150. 2012). The deposit feeding behavior of both 5. BOESE, B. L., M. WINSOR, H. LEE, S. M. nasuta and M. inquinata showed varia- ECHOLS, J. PELLETIER, and R. RAN- tion in response to water flow. Their inhalant DALL. 1995. PCB cogeners and hexachlo- siphons extended farther, allowing for de- robenzene biota sediment accumulation posit feeding over a larger area, when water factors for Macoma nasuta exposed to flow was lower (Levinton 1991). sediments with different total organic car- Predators: Small clams are fed upon by bon contents. Environmental Toxicology crabs (e.g., Cancer productus), seastars and Chemistry. 14:303-310. (e.g., Pisaster spp.), as well as the snail 6. BRINK, L. A. 2001. Mollusca: Bivalvia, p. Polinices lewisii (Haderlie and Abbott 1980; 129-149. In: Identification guide to larval

marine invertebrates of the Pacific North- Journal of Experimental Marine Biology west. A. Shanks (ed.). Oregon State Uni- and Ecology. 175:253-274. versity Press, Corvallis, OR. 16. GALLUCCI, V. F., and J. HYLLEBERG. 7. BRUSCA, G. J., and R. C. BRUSCA. 1976. A quantification of some aspects of 1978. A naturalist's seashore guide. Mad growth in the deposit feeding bivalve Ma- River Press, Arcata, CA. coma nasuta. Veliger. 19:59-67. 8. CADDY, J. F. 1967. Maturation of gam- 17. HADERLIE, E. C., and D. P. ABBOTT. etes and spawning in Macoma balthica 1980. Bivalvia: the clams and allies, p. 355 (L.). Canadian Journal of Zoology. -410. In: Intertidal invertebrates of Califor- 45:955-965. nia. R. H. Morris, D. P. Abbott, and E. C. 9. —. 1969. Development of mantle organs, Haderlie (eds.). Stanford University Press, feeding, and locomotion in postlarval Ma- California. coma balthica (L.) (Lamellibranchiata). 18. HYLLEBERG, J., and V. F. GALLUCCI. Canadian Journal of Zoology. 47:609- 1975. Selectivity in feeding by deposit- 617. feeding bivalve Macoma nasuta. Marine 10. CHO, Y., D. W. SMITHENRY, U. Biology. 32:167-178. GHOSH, A. J. KENNEDY, R. N. MILL- 19. KABAT, A. R., and D. O'FOIGHIL. 1987. WARD, T. S. BRIDGES, and R. G. LU- Phylum Mollusca, Class Bivalvia, p. 309- THY. 2007. Field methods for amending 353. In: Reproduction and development of marine sediment with activated carbon marine invertebrates of the northern Pacif- and assessing treatment effectiveness. ic Coast. M. F. Strathmann (ed.). Universi- Marine Environmental Research. 64:541- ty of Washington Press, Seattle, WA. 555. 20. KEEN, A. M., and E. COAN. 1974. Marine 11. COAN, E. V. 1971. The Northwest Amer- molluscan genera of western North Ameri- ican Tellinidae. California Malacozoologi- ca: an illustrated key. Stanford University cal Society, Berkeley. Press, Stanford, CA. 12. COAN, E. V., and P. VALENTICH- 21. KOZLOFF, E. N. 1974. Seashore life of SCOTT. 2007. Bivalvia, p. 807-859. In: Puget Sound, the Strait of Georgia, and The Light and Smith manual: intertidal the San Juan Archipelago and adjacent invertebrates from central California to regions. University of Washington Press, Oregon. J. T. Carlton (ed.). University of Seattle and London. California Press, Berkeley, CA. 22. —. 1991. Malacobdella siliquae sp. nov. 13. CROOKS, J. A. 2001. Assessing invader and Malacobdella macomae sp. nov., roles within changing ecosystems: His- commensal nemerteans from bivalve mol- torical and experimental perspectives on lusks on the Pacific Coast of North Ameri- an exotic mussel in an urbanized lagoon. ca. Canadian Journal of Zoology. 69:1612- Biological Invasions. 3:23-36. 1618. 14. DUNNILL, R. M., and D. V. ELLIS. 1969. 23. —. 1993. Seashore life of the northern Pa- Recent species of the genus Macoma cific coast: an illustrated guide to northern (Pelecypoda) in British Columbia. Nation- California, Oregon, Washington, and Brit- al Museum of Canada, National Histori- ish Columbia. University of Washington cal Papers. 45:1-34. Press, Seattle. 15. EVERETT, R. A. 1994. Macroalgae in 24. LAMMENS, J. J. 1967. Growth and repro- marine soft-sediment communities: ef- duction in a tidal flat population of Macoma fects on benthic faunal assemblages. balthica (L.). Netherlands Journal of Sea

Research. 3:315-382. 35. SCHELL, S. C. 1989. The structure of the 25. LEVINTON, J. S. 1991. Variable feeding nervous system of Graffilla pugetensis, a behavior in three species of Macoma parasite in the pericardial cavity of the (Bivalvia, Tellinacea) as a response to bent-nose clam, Macoma nasuta. Journal water flow and sediment transport. Ma- of Parasitology. 75:428-430. rine Biology. 110:375-383. 36. SCOTT, P. V., and J. A. BLAKE. 1998. 26. LUTTIKHUIZEN, P. C., J. DRENT, K. T. The Mollusca Part 1: the Aplacophora, C. A. PEIJNENBURG, H. W. VAN DER Polyplacophora, Scaphopoda, Bivalvia VEER, and K. JOHANNESSON. 2012. and Cephalopoda. Taxonomic atlas of the Genetic architecture in a marine hybrid benthic fauna of the Santa Maria Basin zone: comparing outlier detection and and the Western Santa Barbara Channel. genomic clines analysis in the bivalve Vol. 8. Santa Barbara Museum of Natural Macoma balthica. Molecular Ecology. History, Santa Barbara, CA. 21:3048-3061. 37. TURGEON, D. D., J. F. QUINN, A. E. BO- 27. MACGINITIE, G. E., and N. MACGINI- GAN, E. V. COAN, F. G. HOCHBERG, W. TIE. 1949. Natural history of marine ani- G. LYONS, P. M. MIKKELSEN, R. J. mals. McGraw-Hill Book Co., New York. NEVES, C. F. E. ROPER, G. ROSEN- 28. MARRIAGE, L. D. 1954. The bay clams BERG, B. ROTH, A. SCHELTEMA, F. G. of Oregon. Contribution No. 20. Fish THOMPSON, M. VECCHIONE, and J. D. Commission of Oregon, s.l. WILLIAMS. 1998. Common and scientific 29. PACKARD, E. L. 1918. Molluscan fauna names of aquatic invertebrates from the from San Francisco Bay. Zoology. United States and Canada: mollusks. 14:199-452. American Fisheries Society Special Publi- 30. PRUELL, R. J., N. I. RUBINSTEIN, B. K. cation 26. American Fisheries Society, Be- TAPLIN, J. A. LIVOLSI, and R. D. BOW- thesda, MD. EN. 1993. Accumulation of polychlorinat- 38. VOLKENBORN, N., C. MEILE, L. POL- ed organic contaminants from sediment ERECKY, C. A. PILDITCH, A. NORKKO, by three benthic marine species. Ar- J. NORKKO, J. E. HEWITT, S. F. chives of Environmental Contamination THRUSH, D. S. WETHEY, and S. A. and Toxicology. 24:290-297. WOODIN. 2012. Intermittent bioirrigation 31. RAE, J. G. 1978. Reproduction in two and oxygen dynamics in permeable sedi- sympatric species of Macoma (Bivalvia). ments: An experimental and modeling Biological Bulletin. 155:207-219. study of three tellinid bivalves. Journal of 32. —. 1979. Population dynamics of two Marine Research. 70:794-823. sympatric species of Macoma (Mollusca, 39. WERNER, I., S. J. TEH, S. DATTA, X. Q. Bivalvia). Veliger. 21:384-399. LU, and T. M. YOUNG. 2004. Biomarker 33. RICKETTS, E. F., and J. CALVIN. 1971. responses in Macoma nasuta (Bivalvia) Between Pacific tides. Stanford Universi- exposed to sediments from northern San ty Press, Stanford, California. Francisco Bay. Marine Environmental Re- 34. ROE, P., J. L. NORENBURG, and S. search. 58:299-304. MASLAKOVA. 2007. Nemertea, p. 221- 40. WINSOR, M. H., B. L. BOESE, H. LEE, R. 233. In: Light and Smith manual: intertid- C. RANDALL, and D. T. SPECHT. 1990. al invertebrates from central California to Determination of the ventilation rates of Oregon. J. T. Carlton (ed.). University of interstitial and overlying water by the clam California Press, Berkeley, CA. Macoma nasuta. Environmental Toxicolo-

gy and Chemistry. 9:209-213. 41. ZONNEVELD, J., M. K. GINGRAS, C. A. HODGSON, L. P. MCHUGH, R. A. MY- ERS, J. A. SCHOENGUT, and B. WET- THUHN. 2014. Biotic segregation in an upper mesotidal dissipative ridge and runnel succession, West Salish Sea, Vancouver Island, British Columbia, p. 169-194. In: Experimental approaches to understanding fossil organisms: lessons from the living. Vol. 41. D. I. Hembree, B. F. Platt, and J. J. Smith (eds.). Updated 2015 T.C. Hiebert