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Phylum: Arthropoda, Crustacea producta Class: Order: A Section: Brachyura Family:

Taxonomy: was originally the mouth and extend to cover the mandibles described as Epialtus productus by Randall in (Ruppert et al. 2004). 1840, but was later moved to the Carapace: Carapace remarkably Pugettia. Current synonyms for P. producta smooth and mature specimens are practically include E. products, and P. productus (Ng et hairless (Garth 1958). Sides of carapace are al. 2008). almost parallel with prominent posterolateral teeth (Pugettia, Carlton and Kuris 1975). Description Posterior margin convex medially (Garth Size: Pugettia producta is the largest of the 1958) and carapace decidedly longer than kelp (Wicksten 2011) and the largest wide (Wicksten 2011). individual, on record, was 93 mm in width and Frontal Area: Rostrum deeply 107 mm in length. Oregon specimens are notched and bifid, small and with horns larger than those in southern California. The bearing hooked setae (Garth 1958; Wicksten average male carapace is 71 mm in length 2011) (Figs. 1, 2). and 62 mm in width, female carapace is 69 Teeth: Dorsal surface smooth with mm in length and 59 mm in width (Wicksten small pre- and post-orbital teeth (Fig. 2). 2011). Large anterolateral (hepatic) teeth. Large Color: Body color highly dependent on hepatic tooth is joined to postorbital tooth. A season, molting stage and ingested large tooth is also present between macroalgal pigments (Hultgren and anterolateral tooth and posterior margin. Stachowicz 2008) and ranges from light olive Pereopods: Almost cylindrical green to dark brown or black. Ventrally, (Rathbun 1925) and decreasing in length females and juveniles are often yellow and posteriorly with slender dactyls that bear males bright red (see Wicksten 2011). spinules (Fig. 1) (Garth 1958; Wicksten 2011). General Morphology: The body of decapod Legs shorter and stouter than in other can be divided into the Pugettia . cephalothorax (fused head and thorax) and Chelipeds: Large and well abdomen. They have a large plate-like developed, especially in mature males, where carapace dorsally, beneath which are five they are stout and shorter than the first pairs of thoracic appendages (see chelipeds walking legs. The hand long and narrow and pereopods) and three pairs of (sometimes inflated), fingers (dactyls) are maxillipeds (see mouthparts). The abdomen slender, pointed posteriorly and curved and associated appendages are reduced and inward, inner margins are dentate and often folded ventrally (Decapoda, Kuris et al. 2007). gaping (males). Female chelipeds, on the Cephalothorax: other hand, are slender (Fig. 3) (Rathbun Eyes: Eyes small and distance 1925). Female cheliped more slender than between them less than one third carapace male (Wicksten 2011). width (adults). Abdomen (Pleon): Abdomen narrow in Antennae: male, broad in female (Fig. 4). Consists of Mouthparts: The mouth of decapod seven segments (Fig. 4). crustaceans comprises six pairs of Telson & Uropods: appendages including one pair of mandibles Sexual Dimorphism: Male and female (on either side of the mouth), two pairs of brachyuran crabs are easily differentiable. maxillae and three pairs of maxillipeds. The The most conspicuous feature, the abdomen, maxillae and maxillipeds attach posterior to is narrow and triangular in males while it is

Hiebert, T.C. and A. Burgess. 2015. Pugettia producta. 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/1794/12731 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to [email protected] wide and flap-like in females. Additionally, (Garth and Abbott 1980; Coe 1902; Kuris et males have one large chela (see Chelipeds) al. 2007). and two pleopod pairs specialized for Abundance: Most common kelp crab in copulation however, the third and fourth Coos Bay estuary. pleopods are absent. Females, on the other hand, have all four pleopod pairs, each with Life-History Information long setae for egg attachment (Brachyura, Reproduction: All decapod Kuris et al. 2007). females attach recently laid gelatinous egg masses to their pleopods. The outer embryo Possible Misidentifications membrane thickens and a strand develops Among the Epialtidae, there are four local that attaches each embryo to pleopod setae species, three of which belong to the genus (Decapoda, Kuris et al. 2007). Ovigerous Pugettia (Kuris et al. 2007). Besides P. females have been observed year-round in producta, and Pugettia richii Monterey Bay, California, although rarely are two smaller species found in the during some months. In Puget Sound, northwest. Both are smaller and have a Washington largest broods are observed in greater distance between the eyes (about half November to January. Brood numbers range the carapace width) than P. producta. from 34,000 to 84,000 with an average of Additionally, both have tuberculate carapace 61,000 embryos that are yellow early in surfaces, constrictions between the hepatic development and become orange-red to and branchial teeth and long walking legs brown at more advanced stages (Jaffe et al. (Garth 1958, Wicksten 2011). Neither P. 1987). Developmental timelines are variable gracilis nor P. richii have the smooth surface with latitude from 28–31 days in Monterey, or straight carapace sides of P. producta. California to almost a year in Puget Sound, Pugettia gracilis can be similar in color to P. Washington (Knudsen 1964; Jaffe et al. producta, but P. richii is usually red with legs 1987). Females produce broods regularly banded (Wicksten 2011). Members of the and some authors suggest as often as local epialtid genus Mimulus lack monthly (Hines 1981; Jaffe et al. 1987). posterolateral spines. Carcinonemertes epialti is a nemertean predator of P. producta eggs (Coe 1902). Up Ecological Information to 100 nemerteans were found on a single Range: Type locality is northern California. crab (Monterey, California, Coe 1902). The Known range includes Prince of Wales Island, reproduction and life-cycle of C. epialti is Alaska to Point Asuncion, Baja California dependent upon and corresponds to that of its (Garth 1958; Wicksten 2011). host species. However, this nemertean is not Local Distribution: Coos Bay distribution in host specific (unlike Carcinonemertes errans South Slough. Oregon distribution includes on Cancer magister) and occurs amongst egg various protected outer shores and estuaries. masses of other species including Habitat: Off hard substrate and amongst Hemigrapsus oregonensis, H. nudus, eelgrass and kelp (Egregia, Ricketts and Pachygrapsus crassipes (Roe et al. 2007). Calvin 1971). Also occurs in tidepools on Interestingly, it may be more common on H. , on pilings in bays and in oregonensis than P. producta, for which it Enteromorpha, but prefers (juveniles) was described (Kuris 1993; Kuris et al. 2007). (Garth and Abbott 1980). Larva: The larval development of P. product Salinity: Collected at salinities of 30. Does has not been described in detail. not osmoregulate or tolerate brackish water Development consists of zoea (two) and (Garth and Abbott 1980). megalopa stages (Hines 1981). The zoea of Temperature: members of the family Majidae (now Tidal Level: Intertidal to 80 meters (Garth Epialtidae) can be recognized by the 1958; Wicksten 2011). presence of a rostral spine only, lack of lateral Associates: Sometimes hosts parasitic spines and an antenna protopod that is nearly , Sacculina. Eggs parasitized by equal in length to the rostrum. The carapace nemertean worm Carcinonemertes epialti of megalopae are 1.6 mm in length and 1.2

Hiebert, T.C. and A. Burgess. 2015. Pugettia producta. 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. mm in width and is larger than its congener, 2. COE, W. R. 1902. The nemertean P. gracilis (Puls 2001). Recruitment of newly parasites of crabs. American metamorphosed individuals is mostly in Naturalist:431-450. intertidal and shallow subtidal eelgrass and 3. GARTH, J. S. 1958. Brachyura of the surfgrass zones (Jaffe et al. 1987). Pacific coast of America: Oxyrhyncha. Juvenile: Juveniles (about 3 mm long) can Allan Hancock Pacific Expedition. be constricted at the sides like P. richii (Garth 21:188-193. 1958). Newly settled individuals bear tufts of 4. GARTH, J. S., and D. P. ABBOTT. setae on the lateral margins of the carapace 1980. Brachyura: The true crabs, p. and are yellow in color, ventrally (Wicksten 594-630. In: Intertidal invertebrates of 2011). California. R. H. Morris, D. P. Abbott, Longevity: and E. C. Haderlie (eds.). Stanford Growth Rate: Growth occurs in conjunction University Press, Stanford, CA. with molting. In pre-molting periods the 5. GROSSMAN, G. D. 1986. Food epidermis separates from the old cuticle and resource partitioning in a rocky a dramatic increase in epidermal cell growth intertidal assemblage. Journal of occurs. Post-molt individuals will have soft Zoology Series B. 1:317-356. shells until a thin membranous layer is 6. HINES, A. H. 1981. Life history deposited and the cuticle gradually strategies of spider crab (Majidae). hardens. During a molt decapods have the American Zoologist. 21:990-990. ability to regenerate limbs that were 7. —. 1982. Coexistence in a : previously autotomized (Kuris et al. 2007). size, population dynamics, and Food: Pugettia producta is primarily resource partitioning in a guild of herbivorous, preferring brown algae, but will spider crabs (Brachyura: Majidae). eat , and fish pieces in the Ecological Monographs. 52:179-198. laboratory (Knudsen 1964). Ontogenetic 8. HULTGREN, K. M., and J. J. changes from red intertidal algae to brown STACHOWICZ. 2008. Alternative algae (kelp) has been reported by Hines camouflage strategies mediate (1982). risk among closely related Predators: (especially juveniles), co-occurring kelp crabs. Oecologia. larger crabs and sea otters (Grossman 1986; 155:519-528. Hultgren and Stachowizc 2008). Adult P. 9. JAFFE, L. A., C. F. NYBLADE, R. B. producta are aggressive with a strong pinch FORWARD, and S. SULKIN. 1987. and thus, few predators. Additionally, color Phylum or subphylum Crustacea, camouflage effective in reducing predation class Malacostraca, order Decapoda, (Hultgren and Stachowicz 2008). Brachyura, p. 451-475. In: Behavior: A nocturnal feeder, P. producta is Reproduction and development of an active species, particularly those in rocky marine invertebrates of the northern tide pools (Rathbun 1925). Individuals Pacific coast. M. F. Strathmann (ed.). occasionally attach pieces of algae, which will University of Washington Press, be consumed later, to their rostrum (Wicksten Seattle, WA. 2011). 10. KNUDSEN, J. W. 1964. Observations of the reproductive cycles and ecology Bibliography of the common Brachyura and crablike Anomura of Puget Sound, 1. CARLTON, J. T., and A. M. KURIS. Washington. Pacific Science. 18:3-33. 1975. Keys to decapod crustacea, p. 11. KURIS, A. M. 1993. Life cycles of 385-412. In: Light's manual: intertidal nemerteans that are symbiotic egg invertebrates of the central California predators of decapod crustacea: coast. S. F. Light, R. I. Smith, and J. T. to host life histories. Carlton (eds.). University of California Hydrobiologia. 266:1-14. Press, Berkeley. 12. KURIS, A. M., P. S. SADEGHIAN, J. T. CARLTON, and E. CAMPOS. 2007.

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12731 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to [email protected] Decapoda, p. 632-656. In: The Light and Smith manual: intertidal invertebrates from central California to Oregon. J. T. Carlton (ed.). University of California Press, Berkeley, CA. 13. NG, P. K. L., D. GUINOT, and P. J. F. DAVIE. 2008. Systema brachyurorum: Part I. Annotated checklist of the extant Brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement. 17:1-286. 14. PULS, A. L. 2001. Arthropoda: Decapoda, p. 179-250. In: Identification guide to larval marine invertebrates of the Pacific Northwest. A. Shanks (ed.). Oregon State

University Press, Corvallis, OR. 15. RATHBUN, M. J. 1925. The Spider

crabs of America. Bulletin of the United States Natural Museum. 129:167-172. 16. RICKETTS, E. F., and J. CALVIN. 1971. Between Pacific tides. Stanford University Press, Stanford, California. 17. ROE, P., J. L. NORENBURG, and S. MASLAKOVA. 2007. Nemertea, p. 221-233. In: Light and Smith manual: intertidal invertebrates from central California to Oregon. J. T. Carlton (ed.). University of California Press, Berkeley, CA. 18. RUPPERT, E. E., R. S. FOX, and R. D. BARNES. 2004. Invertebrate zoology: a functional evolutionary approach. Thomson Brooks/Cole, Belmont, CA. 19. WICKSTEN, M. K. 2011. Decapod crustacea of the Californian and Oregonian Zoogeographic Provinces. http://escholarship.org/uc/it em/7sk9t2dz. Scripps Institution of Oceanography, UC San Diego, San Diego, CA.

Hiebert, T.C. and A. Burgess. 2015. Pugettia producta. 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.