Phylum:

Nucella lamellosa Class: ,

Order: A gammarid amphipod Family: Muricoidea, ,

Taxonomy: was previously called 1974); encrusted, smooth. Thais. Thais is now reserved for subtropical Suture: Impressed, distinct, but not a and tropical . For a more detailed deep groove. review of gastropod , see Keen Anterior (Siphonal) Canal: Short, but and Coan (1974) and McLean (2007). longer than other Nucella species; narrow, slot-like, not spout-like (i.e. with edges touch- Description ing, making a closed tube: see Possible Misi- Size: To 50 mm in California (Abbott and dentifications). Not separated from large Haderlie 1980), 100 mm Puget Sound and whorl by revolving groove (fig. 1). north (Kozloff 1974); largest specimen illus- Umbilicus: Small, often closed (fig. 1). trated, 54 mm (fig. 1). Largest of the Nucella Aperture: Almost 1/2 length shell; . ovate to quadrate in outline, with a siphonal Color: White to brown, some are pink, lav- notch, but no anal notch (fig. 1). Widest part ender or orange tan; not highly polished. In- of aperture (generally near its middle) at least side whitish, sometimes with color showing half as wide as shell (Kozloff 1974). through. Outer Lip: Thickened, smooth, without General Morphology: denticles on posterior portion of aperture Shell: (near anal notch) no single strong tooth on Shape: Shell heavy, solid, strong; edge near anterior canal (see Possible Misi- spirally coiled, fusiform (spindle-shaped). 5- dentifications). Outer lips rounding smoothly 7 whorls; nuclear whorl small, inconspicu- to anterior end of shell. At least one row of ous. Spire usually high; siphonal canal rela- denticles within lip (fig. 1). tively long for genus; aperture ovate, almost Operculum: Usually large enough to 1/2 shell length. close aperture; conspicuous, with strong spi- Sculpture: Extremely variable. Spire ral lines; with nucleus on one side (fig. la). and base have similar sculpture: genus Nu- Eggs: Vase-shaped, yellow, about 10 mm cella (McLean 2007, Keen and Coan 1974). long; in clusters on underside of rocks (Abbott Axial ribs present (fig. 1). Three chief varia- and Haderlie 1980); called "sea "; (fig. tions with many gradations: lamellar variety 1B). with strong axial ribs, developed in quiet wa- ter specimens into frilly ruffles (fig. 4); Nucel- Possible Misidentifications la from rough conditions are smooth, with Nucella can be distinguished from oth- only faint axial sculpture (figs. 1, 3); and er predatory estuarine snails by its sculpture, strongly sculptured spirally with one to two which is the same on the whorls and spire, by strong horizontal ribs at top of each whorl the large last whorl and by the ovate aperture and smaller ribs below; axial sculpture only (about 1/2 the shell length). Unlike Nassarius, between ribs. This third variety has flattened it has no distinct revolving furrow setting off and angled whorls (fig. 2) (Kozloff 1974). the body whorl from the anterior canal (Keen Columella: Without folds (Kozloff and Coan 1974). It has no single strong tooth

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]

Bering, N., T. Hext and E. Parker. 2017. . 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 Biol- ogy, Charleston, OR.

on the anterior margin of the outer lip, as in concentrations. . There are no columellar folds as Temperature: Cold to temperate waters. Ge- in Olivella, Buccinus, etc. The siphonal canal ographic distribution would indicate a prefer- is not spout-like, as in , and ence for cool temperatures. Lower part of bay . does not generally have high temperatures. There are several species of Nucella Smallest individuals have highest thermal lim- in the northwest: its; snails active at 0-30 °C (Bertness 1977). , the file dogwhelk, is a Tidal Level: Found at low intertidal, lower subtidal snail with about 16 alternating large than other species of the genus. Largest ani- and small file-like spiral ridges on the large mals lowest in tidal range (Bertness 1977). whorl. It is rare, whitish to brown in color, Associates: Primary prey is short-spired and somewhat smaller than N. Balanus, which shows reduced settlement lamellosa (to 43 mm). and metamorphosis in areas occupied or pre- , the channeled viously occupied by N. lamellosa (Johnson dogwhelk, is white to or orange, sometimes and Strathmann 1989); found with porcelain banded. It has a high spire, a prominent crab Petrolisthes, brachyuran crabs Hemi- shoulder below the deep suture, and round- grapsus and Cancer oregonensis, chiton Mo- ed spiral ridges of equal size with axial la- palia, isopod Idotea, anemones Anthopleura mellae between them. It is small, to just over elegantissima and A. artemesia, nudibranch 30 mm. Usually found in beds, it is Onchidoris, gastropods Tegula and Pisaster rare in bays (Kozloff 1974). ochraceus. Discarded N. lamellosa shells of- is the other Nucella ten inhabited by the hermit crab Pagurus hir- most often found in estuaries; it usually oc- suitusculus. curs in heavier surf than N. lamellosa. Weight: Largest collected (including shell) 28 Called the rock-dwelling dogwinkle, it is gen- gr. (wet). erally only up to 20 mm long. This snail has Abundance: One of the most abundant inter- alternately large and small, often nodulose, tidal snails of the northwest; becomes less spiral ridges over most of the shell. (These abundant in California. By far the most com- ridges are often obscure). It has no noticea- mon Nucella species in the Coos Bay estuary. ble axial sculpture. Found in the mid- and Life-History Information high intertidal in mussel beds, N. ostrina is Reproduction: Mates in winter and spring easily confused with variation of N. lamel- (California) by aggregations of snails; individ- losa (fig. 2). uals become sexually mature in 4th year, Ecological Information when they often return to their hatching site Range: Bering Strait to central California and join a breeding group (Abbott and Hader- (Abbott and Haderlie 1980). lie 1980); individuals tend to breed with same Local Distribution: Coos Bay: Pigeon group. Gestation is approximately 20 months. Point, Empire; Umpqua estuary: Ziolkouski Spawning occurs in June. Egg capsules de- Beach (1/2 mile from mouth). posited synchronously with other females; de- Habitat: On rocks with mud, sand substrate; velopment varies with temperature: snails often in protected bays (McLean 2007); be- emerge after 140 days (at 6.8°C), after 67-91 low mussel beds on outer shores. days (9.6-11 °C) (Seavy 1977). Capsules pro- Salinity: Collected at 30: lower, more ma- tect embryos from low salinity stress by re- rine parts of bays with more constant saline ducing the rate at which the osmotic concen-

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]

tration of intracapsular fluid decreases of a known predator (Chattopadhyay and (Pechenik 1982). Capsules rarely contain Baumiller 2007). "nurse eggs" (sterile eggs to be consumed Bibliography by the developing snail larvae): nearly all eggs are fertile. Just over half of eggs reach 1. ABBOTT, D.P. and E. C. HADERLIE. hatching stage. There is high mortality 1980. Prosobranchia: marine snails, p.230 among young snails and of 1000 eggs (from -307. In: Intertidal invertebrates of Califor- one female, one year) probably fewer than nia. R.H. MORRIS, D. P. ABBOTT, and E. 10 grow to one year of age (Lyons and C. HADERLIE (eds.). Stanford University Spight 1973). Press, Stanford, California. Larva: 2. APPLETON, R. D., and A.R. PALMER. Juvenile: 1988. Water-borne stimuli released by Longevity: Sexually mature at four years predatory crabs and damaged prey induce (Abbott and Haderlie 1980). more predator-resistant shells in a marine Growth Rate: Varies greatly with food sup- gastropod. Proceedings of the National ply. Shell growth, type, dependent on food: Academy of Sciences. 85:4387-4391. barnacle diet produces heavy, stout shells. 3. BERTNESS, M. D. 1977. Behavioral and Water-soluble chemical cues released by ecological aspects of shore-level gradients and by damaged conspe- in Thais lamellosa and Thais emarginata. cifics induced N. lamellosa to improve the Ecology. 58:86-97. defense effectiveness (thickness and mor- 4. CHATTOPADHYAY, D., and T.K. BAU- phology) of their shells (Appleton and Palm- MILLER. 2007. Drilling under threat: An er 1988). experimental assessment of the drilling Food: Primarily : Balanus glandula behavior of Nucella lamellosa in the pres- and B. cariosus, on which it is the primary ence of a predator. Journal of Experi- predator (Puget Sound) (Kozloff 1974). Mus- mental Marine Biology and Ecology. sels (outer shores), periwinkles and other 352:257-266. mollusks. penetrates shell of prey 5. JOHNSON, L. E., and R. R. STRATH- with aid of secretions from boring organ on MANN. 1989. Settling barnacle larvae foot (Abbott and Haderlie 1980). avoid substrata previously occupied by a Predators: Crabs (Cancer productus, Hemi- mobile predator. Journal of Experimental grapsus oregonensis) egg capsules and Marine Biology and Ecology. 128:87–103. young snails heavily preyed upon by other 6. KEEN, A. M. and E.V. COAN. 1974. Ma- Nucella. Nucella lamellosa exhibits hatching rine molluscan genera of Western North plasticity by taking longer to hatch in the America; an illustrated key. 2d ed. Stan- presence of predators (Hemigrapsus ore- ford, Calif.: Stanford University Press. gonensis) and increasing rate of hatching in 7. KOZLOFF, E.N. 1974. Keys to the marine the presence of conspecific adults (Miner et invertebrates of Puget Sound, the San al. 2010). Juan Archipelago, and adjacent regions. Behavior: The presence of a natural preda- University of Washington Press, Seattle & tor (Cancer gracilis) while N. lamellosa is London. feeding on Mytilus californianus results in an 8. LYONS, A., and T. M. SPIGHT. 1973. Di- increased number of incomplete drill holes, versity of feeding mechanisms among em- suggesting that N. lamellosa abandons its bryos of Pacific Northwest Thais. The Veli- prey more frequently when in the presence ger. 16:189-194.

Bering, N., T. Hext and E. Parker. 2017. Nucella lamellosa. 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 Biol- ogy, Charleston, OR.

9. MCLEAN, J.H. 2007. Shelled gastropo- da, p. 713-753. In: The Light and Smith manual: intertidal invertebrates from cen- tral California to Oregon. J. T. Carlton (ed.). University of California Press, Berkeley. 10. MINER, B. G., A.D. DONOVAN, and K.E. ANDREWS. 2010. Should I stay or should I go: predator and conspecific- induced hatching in a marine snail. Oecologia. 163:69-78. 11. PECHENIK, J. A. 1982. Ability of some gastropod egg capsules to protect against low-salinity stress. Journal of Ex- perimental Marine Biology and Ecology. 63:195–208. 12. SEAVY, D. K. 1977. Seasonal gameto- genesis and egg laying in the proso- branch gastropods Nucella lamellosa, , Searlesia dira and columbiana on the Oregon coast. Ph.D. Oregon State University. Updated 2017 N. Bering, E. Parker, and T. Hext

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]