Phylum: lamellosa Class: , Prosobranchia Order: The wrinkled or frilled dogwinkle Family:

Taxonomy: Nucella was previously called groove. . Thais is now reserved for subtropical Anterior (Siphonal) Canal: Short, but longer and tropical . For a more detailed than other Nucella species; narrow, slot-like, review of gastropod , see Keen and not spout-like (i.e. with edges touching, Coan (1974) and McLean (2007). making a closed tube: see Possible Misidentifications). Not separated from large Description by revolving groove (fig. 1). Size: To 50 mm in California (Abbott and : Almost 1/2 length shell; ovate to Haderlie 1980), 100 mm Puget Sound and quadrate in outline, with a siphonal notch, but north (Kozloff 1974); largest specimen no anal notch (fig. 1). Widest part of aperture illustrated, 54 mm (fig. 1). Largest of the (generally near its middle) at least half as Nucella . wide as shell (Kozloff 1974). Color: White to brown, some are pink, : Small, often closed (fig. 1). lavender or orange tan; not highly polished. : Usually large enough to close Inside whitish, sometimes with color showing aperture; conspicuous, with strong spiral through. lines; with nucleus on one side (fig. la). Shell Shape: Shell heavy, solid, strong; Eggs: Vase-shaped, yellow, about 10 mm spirally coiled, fusiform (spindle-shaped). 5-7 long; in clusters on underside of rocks (Abbott whorls; nuclear whorl small, inconspicuous. and Haderlie 1980); called "sea "; (fig. usually high; relatively 1B). long for genus; aperture ovate, almost 1/2 shell length. Possible Misidentifications : Extremely variable. Spire and Nucella can be distinguished from other base have similar sculpture: genus Nucella predatory estuarine snails by its sculpture, (McLean 2007, Keen and Coan 1974). Axial which is the same on the whorls and spire, by ribs present (fig. 1). Three chief variations the large last whorl and by the ovate aperture with many gradations: lamellar variety with (about 1/2 the shell length). Unlike , strong axial ribs, developed in quiet water it has no distinct revolving furrow setting off specimens into frilly ruffles (fig. 4); Nucella the from the anterior canal (Keen from rough conditions are smooth, with only and Coan 1974). It has no single strong tooth faint axial sculpture (figs. 1, 3); and strongly on the anterior margin of the outer , as in sculptured spirally with one to two strong Acanthina. There are no columellar folds as in horizontal ribs at top of each whorl and Olivella, Buccinus, etc. The siphonal canal is smaller ribs below; axial sculpture only not spout-like, as in Ocenebra, and between ribs. This third variety has flattened Ceratostoma.

and angled whorls (fig. 2) (Kozloff 1974). There are several species of Nucella in the Outer Lip: Thickened, smooth, without northwest: denticles on posterior portion of aperture , the file dogwhelk, is a (near anal notch) no single strong tooth on subtidal snail with about 16 alternating large edge near anterior canal (see Possible and small file-like spiral ridges on the large Misidentifications). Outer lips rounding whorl. It is rare, whitish to brown in color, smoothly to anterior end of shell. At least one short-spired and somewhat smaller than N. row of denticles within lip (fig. 1). lamellosa (to 43 mm). : Without folds (Kozloff 1974); , the channeled encrusted, smooth. dogwhelk, is white to or orange, sometimes : Impressed, distinct, but not a deep banded. It has a high spire, a prominent

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 Biology, Charleston, OR.

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12913 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected]

shoulder below the deep suture, and rounded Weight: Largest collected (including shell) 28 spiral ridges of equal size with axial lamellae gr. (wet). between them. It is small, to just over 30 mm. Abundance: One of the most abundant Usually found in beds, it is rare in intertidal snails of the northwest; becomes bays (Kozloff 1974). less abundant in California. By far the most is the other Nucella most common Nucella species in the Coos Bay often found in estuaries; it usually occurs in estuary. heavier surf than N. lamellosa. Called the rock-dwelling dogwinkle, it is generally only Life History Information up to 20 mm long. This snail has alternately Reproduction: Mates in winter and spring large and small, often nodulose, spiral ridges (California) by aggregations of snails; over most of the shell. (These ridges are often individuals become sexually mature in 4th obscure). It has no noticeable axial sculpture. year, when they often return to their hatching Found in the mid- and high intertidal in site and join a breeding group (Abbott and mussel beds, N. ostrina is easily confused Haderlie 1980); individuals tend to breed with with variation of N. lamellosa (fig. 2). same group. Gestation is approximately 20 months. Spawning occurs in June. Egg Ecological Information capsules deposited synchronously with other Range: Bering Strait to central California females; development varies with (Abbott and Haderlie 1980). temperature: snails emerge after 140 days (at Local Distribution: Coos Bay: Pigeon Point, 6.8°C), after 67-91 days (9.6-11 °C) (Seavy Empire; Umpqua estuary: Ziolkouski Beach 1977). Capsules protect embryos from low (1/2 mile from mouth). salinity stress by reducing the rate at which Habitat: On rocks with mud, sand substrate; the osmotic concentration of intracapsular often in protected bays (McLean 2007); below fluid decreases (Pechenik 1982). Capsules mussel beds on outer shores. rarely contain "nurse eggs" (sterile eggs to be Salinity: Collected at 30: lower, more marine consumed by the developing snail larvae): parts of bays with more constant saline nearly all eggs are fertile. Just over half of concentrations. eggs reach hatching stage. There is high Temperature: Cold to temperate waters. mortality among young snails and of 1000 Geographic distribution would indicate a eggs (from one female, one year) probably preference for cool temperatures. Lower part fewer than 10 grow to one year of age (Lyons of bay does not generally have high and Spight 1973). temperatures. Smallest individuals have Growth Rate: Varies greatly with food supply. highest thermal limits; snails active at 0-30 °C Shell growth, type, dependent on food: (Bertness 1977). diet produces heavy, stout shells. Tidal Level: Found at low intertidal, lower Water-soluble chemical cues released by than other species of the genus. Largest and by damaged lowest in tidal range (Bertness 1977). conspecifics induced N. lamellosa to improve Associates: Primary prey is barnacle the defense effectiveness (thickness and Balanus, which shows reduced settlement morphology) of their shells (Appleton and and metamorphosis in areas occupied or Palmer 1988). previously occupied by N. lamellosa (Johnson Longevity: Sexually mature at four years and Strathmann 1989); found with porcelain (Abbott and Haderlie 1980). crab Petrolisthes, brachyuran crabs Food: Primarily : Balanus glandula Hemigrapsus and Cancer oregonensis, chiton and B. cariosus, on which it is the primary Mopalia, isopod Idotea, anemones predator (Puget Sound) (Kozloff 1974). Anthopleura elegantissima and A. artemesia, (outer shores), periwinkles and other nudibranch Onchidoris, gastropods Tegula mollusks. penetrates shell of prey with and . Discarded N. aid of secretions from boring organ on foot lamellosa shells often inhabited by the hermit (Abbott and Haderlie 1980). crab Pagurus hirsuitusculus. Predators: Crabs (Cancer productus, Hemigrapsus oregonensis) egg capsules and

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 Biology, Charleston, OR.

young snails heavily preyed upon by other barnacle larvae avoid substrata Nucella. Nucella lamellosa exhibits hatching previously occupied by a mobile plasticity by taking longer to hatch in the predator. Journal of Experimental presence of predators (Hemigrapsus Marine Biology and Ecology. 128:87– oregonensis) and increasing rate of hatching 103. in the presence of conspecific adults (Miner et 6. KEEN, A. M. and E.V. COAN. 1974. al. 2010). Marine molluscan genera of Western Behavior: The presence of a natural predator North America; an illustrated key. 2d (Cancer gracilis) while N. lamellosa is feeding ed. Stanford, Calif.: Stanford on Mytilus californianus results in an University Press. increased number of incomplete drill holes, 7. KOZLOFF, E.N. 1974. Keys to the suggesting that N. lamellosa abandons its marine invertebrates of Puget Sound, prey more frequently when in the presence of the San Juan Archipelago, and a known predator (Chattopadhyay and adjacent regions. University of Baumiller 2007). Washington Press, Seattle & London. 8. LYONS, A., and T. M. SPIGHT. 1973. Diversity of feeding mechanisms among embryos of Pacific Northwest Literature Cited Thais. The . 16:189-194. 1. ABBOTT, D.P. and E. C. HADERLIE. 9. MCLEAN, J.H. 2007. Shelled 1980. Prosobranchia: marine snails, gastropoda, p. 713-753. In: The Light p.230-307. In: Intertidal invertebrates and Smith manual: intertidal of California. R.H. MORRIS, D. P. invertebrates from central California to ABBOTT, and E. C. HADERLIE (eds.). Oregon. J. T. Carlton (ed.). University Stanford University Press, Stanford, of California Press, Berkeley. California. 10. MINER, B. G., A.D. DONOVAN, and 2. APPLETON, R. D., and A.R. K.E. ANDREWS. 2010. Should I stay PALMER. 1988. Water-borne stimuli or should I go: predator and released by predatory crabs and conspecific-induced hatching in a damaged prey induce more predator- marine snail. Oecologia. 163:69-78. resistant shells in a marine gastropod. 11. PECHENIK, J. A. 1982. Ability of Proceedings of the National Academy some gastropod egg capsules to of Sciences. 85:4387-4391. protect against low-salinity stress. 3. BERTNESS, M. D. 1977. Behavioral Journal of Experimental Marine and ecological aspects of shore-level Biology and Ecology. 63:195–208. gradients in Thais lamellosa and Thais 12. SEAVY, D. K. 1977. Seasonal emarginata. Ecology. 58:86-97. gametogenesis and egg laying in the 4. CHATTOPADHYAY, D., and T.K. prosobranch gastropods Nucella BAUMILLER. 2007. Drilling under lamellosa, , threat: An experimental assessment of Searlesia dira and Amphissa the drilling behavior of Nucella columbiana on the Oregon coast. lamellosa in the presence of a Ph.D. Oregon State University. predator. Journal of Experimental Marine Biology and Ecology. 352:257- 266. 5. JOHNSON, L. E., and R. R. STRATHMANN. 1989. Settling

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12913 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected]