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Phylum: Arthropoda, Crustacea

Petrolisthes cinctipes Class: Multicrustacea, , Eumalacostraca

Order: , , , The flat porcelain Family: , Porcellanidae

Taxonomy: cinctipes is and Eyes: The eyestalks of P. cinctipes has been a widely used name for this spe- contain neurosecretory cell bodies (z-organs) cies. There are, however, several junior that regulate regeneration, molting and oocyte synonyms including cinctipes, maturation (Kurup 1964a). Porcellana rupicola, and Petrolisthes rupico- : Very long, and often folded lus (for all synonyms see Haig 1960; Wick- posteriorly over carapace sides (Fig. 1). First sten 2011). (basal) joint of antennal peduncle is short and not reaching upper margin of carapace. Description Mouthparts: The mouth of decapod Size: Individuals up to 24 mm in length comprises six pairs of appendag- (carapace width) (Puls 2001; Wicksten es including one pair of mandibles (on either 2011). The illustrated specimen (from Coos side of the mouth), two pairs of maxillae and Bay) is 14 mm in length and weighs 1.7 g. three pairs of maxillipeds. The maxillae and Color: Dark blue-brown and somewhat iri- maxillipeds attach posterior to the mouth and descent (see Plate 20, Kozloff 1993). An- extend to cover the mandibles (Ruppert et al. tennae dark red, maxillipeds bright red- 2004). Second maxillipeds in P. cinctipes are orange and legs blue banded with white highly developed for filter feeding (see Food) (Schmitt 1921). White comma-like marks with long fine hairs and specialized shape for are sometimes present ventrally and cheli- channeling water currents (Fig. 4). The color peds bear a red spot at dactyl base, while of the palps of maxilliped three are of walking legs have a yellow median band on taxonomic importance: blue in P. eriomerus propodus. Dactyls yellow with narrow brown and orange in P. cinctipes (Kozloff 1993; band. Individuals near molting are blue in Kuris et al. 2007). color (Wicksten 2011). Carapace: Round with carapace front General Morphology: The body of decapod triangulate (Petrolisthes, Haig 1960). Cara- crustaceans can be divided into the cepha- pace surface is finely granulate and not lothorax (fused head and thorax) and abdo- rough. No epibranchial (anterolateral) spines men. They have a large plate-like carapace and epimera and lateral portions of carapace dorsally, beneath which are five pairs of tho- are entire (Figs. 1, 2). Carapace about as racic appendages (see chelipeds and pere- long as wide (Wicksten 2011). opods) and three pairs of maxillipeds (see Frontal Area: Triangular and strongly mouthparts). The body of the Porcellani- deflexed with conspicuous median groove dae is crab-like and convex longitudinally (Fig. 1). with small fifth legs resting on carapace (Fig. Teeth: 1) and the abdomen and associated ap- Pereopods: Waking legs 2–4 with a pendages are reduced and folded ventrally. few coarse spines on dactyl, propodus and The body and chelae of Petrolisthes are flat- carpus, but not on merus (Fig. 1). Merus of tened (Kuris et al. 2007). third leg is inflated and carpus is without setae Cephalothorax:

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual : 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. Petrolisthes cinctipes. 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.

while propodus and dactyl bear setae. Fifth broad, not longer than broad as in Petrolis- legs small, elevated and rest on carapace thes. There are three local species: P. (Figs. 1, 3). holosericus, P. pubescens and P. rudis (Kuris Chelipeds: Equal (or almost), broad et al. 2007). and flattened, not thick and rough Petrolisthes species, on the other (Petrolisthes, Schmitt 1921; Kuris et al. hand, have a flattened body and chelae, che- 2007), covered with fine granules (as in lae of equal size with carpus longer than it is carapace) but without setae. Carpus almost wide. Petrolisthes cinctipes is recognizable invariably 1 1/2 times longer than wide and by characteristics of the cheliped carpus. The anterior and posterior margins converge carpus has a long anterior lobe that extends distally (Schmitt 1921; Kuris et al. 2007) more than 1/4 total carpus length, is smooth (Fig. 1). Posterior margin with ridge of and hairless and with margins that converge tubercles flanked by teeth distally. distally (Kuris et al. 2007). Five Petrolisthes Prominent lobe at inner angle (P. cinctipes, species are reported to occur from central Kuris et al. 2007) (Fig. 1). A short tuft of hair California to Oregon including P. cinctipes, P. between fingers present ventrally, but che- cabrilloi, P. eriomerus, P. manimaculis and P. lae are generally hairless (Figs. 1, 2). Merus rathbunae. with conspicuous lobe on anterior margin. Of those, P. eriomerus is superficially (Wicksten 2011). quite like P. cinctipes (Kozloff 1993). This Abdomen (Pleon): Abdomen symmetrical, crab lives under rocks in gravelly substrates short and permanently folded under thorax. and is a little smaller than P. cinctipes. The Seven abdominal plates (Petrolisthes) (Figs. carpus of the chelipeds in P. eriomerus is 1, 2). twice as long as wide (not 1 1/2 times as long) Telson & Uropods: Seventh plate of telson and the carpus margins are parallel, not con- forms tail fan (Fig. 2). Uropods attached to verging. Also, there is no prominent lobe at abdominal segment five. the inner angle and the carpus has scattered Sexual Dimorphism: Not obvious superfi- tubercules, not a finely granulated surface as cially. Inside telson, males have single pleo- in P. cintctipes. Ventrally, the outer edge of pods on abdominal plate two and females the maxillipeds in P. eriomerus is bright blue, have long, branched pleopods on plates 3–5 not red orange (Kozloff 1993; Kuris et al. (not shown). 2007). These two species exhibit a non- overlapping vertical distribution, where P. eri- Possible Misidentifications omerus occurs in the low intertidal and P. Porcelain (Porcellanidae) are cinctipes is found in the mid to high intertidal flattened dorso-ventrally and are often found (Jensen and Armstrong 1991). in small cracks and crevices. Their third P. cabrilloi, P. manimaculis and P. maxillipeds bear long setae, which they use rathbunae are all reported from California: to filter feed and their fifth walking legs are Petrolisthes cabrilloi from Morro Bay, Califor- modified into brushes for grooming. There nia south to Baja California, Mexico and ap- are two genera of porcelain crabs in our ar- parently replaces populations of P. cinctipes ea, Petrolisthes and . Members south of Point Conception, California; P. of the Pachycheles have a thick, rough body manimaculis from Bodega Bay, California and chelae, chelae are unequal, tuberculate south to Baja California, Mexico; P. rathbunae or granular and hairy, not smooth. Further- from Monterey, California to Isla Guadalupe, more, the carpus of the chela is as long as Mexico (Wicksten 2011).

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]

Ecological Information Found only at shore stations and not by Range: Type locality is erroneously indicat- dredging (San Francisco Bay, Schmitt 1921). ed as in Hawaii, but is likely to be near Mon- Associates: Associates include , tu- terey, California (Wicksten 2011). Known nicates, sponges, nudibranch Onchidoris, range includes British Columbia to Point chiton Mopalia, shore crabs Hemigrapsus, Concepcion, California and also islands offs- Cancer oregonensis, predatory gastropod hore of southern California, and Baja Califor- Nucella, and the sea star Pisaster ochraceus. nia (Haig 1960). Abundance: Very common (Haig 1960) (up Local Distribution: Outer, more marine to 860 individuals per m2, Monterey, Califor- portions of large estuaries. Occurs locally in nia) (Barnard et al. 1980). When found, P. Coos Bay (e.g. Pigeon Point) and in Netarts cinctipes is usually abundant (MacGinitie and Bay. MacGinitie 1949; Kuris et al. 2007). Habitat: Protected, semi-protected rocky Life-History Information coasts under rocks and amongst Reproduction: Females ovigerous every beds (Ricketts and Calvin 1971; Kuris et al. month of the year but April, May, September, 2007). Prefers open shores and clear water October and November (Haig 1960; Barnard (Haig 1960) and is not tolerant of sand and et al. 1980) and evidence shows that multiple silt (Jensen and Armstrong 1991; Wicksten males (1–3) may contribute to each brood 2011). (Toonen 2004). In Coos Bay, March is the Salinity: Collected at salinities of 30. month in which the greatest number of fema- Temperature: A mid to high intertidal spe- les are found with developing young. Eggs P. cinctipes cies, is exposed to a wide range are a little over 800 µm in diameter, deep of temperature (0–32˚C, Stillman and Som- scarlet to maroon when extruded and become ero 2000). Recent research involving physi- brownish red as they advance cal factors associated with climate change developmentally (Gonor and Gonor 1973a; P. cinctipes has used as a model organism Barnard et al. 1980). (e.g. Somero 2010). Stress by thermal vari- Larva: Petrolisthes cinctipes larvae were de- ation, more than other physical factors (e.g. scribed by Gonor and Gonor (1973a, b). De- pH, salinity), negatively effects P. cinctipes velopment proceeds via two zoeal larval stag- (Paganini et al. 2014). However, when ac- es and a filter feeding megalopa, each P. climated for a short period of time (6 hrs), marked by a molt (Puls 2001). cinctipes can increase thermotolerance zoea are recognizable as larval stages by (Ronges et al. 2012). Research involving their elongate anterior and posterior carapace elevated pCO2, salinity and lower pH, all of spines (see Fig. 53.1-3, Harvey et al. 2014; which simulate predicted physical changes Puls 2011; Wicksten 2011) and have been associated with climate change, have fo- described as "preposterous P. cinctipes cused on life-history stages (e.g. unicorns" (Ricketts and Calvin 1971) with a Miller et al. 2014). Long exposure (40 d) to long spine to discourage predators. Other low pH reduced juvenile survival and heart characters of zoeal morphology include a tel- rate. Furthermore, embryonic volumes do son posterior margin that is rounded and with not increase at a normal developmental rate long plumose setae. Pachycheles and Petrol- when exposed to lower pH (Ceballos-Osuna isthes species can be distinguished by the et al. 2013). presence of terminal brushes on telson setae, Tidal Level: Mid and upper tidal levels and in that Pachycheles species have only two almost exclusively littoral (Haig 1960). Hiebert, T.C. 2015. Petrolisthes cinctipes. 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.

and Petrolisthes species have brushes on all detritus from water with fan-like second setae (Puls 2001). The megalopa of Petrol- maxillipeds. Feeding behavior evoked by isthes species have long, slender chelipeds presence of amino acids, sugars (Hartman that are dorso-ventrally flattened (as in and Hartman 1976). Despite their mobility, adults) and P. cinctipes megalopae have a Petrolisthes cinctipes is a gregarious species, cheliped carpus with a single spine on the and increases in conspecific density have inner margin and an inconspicuous central been shown to reduce growth rate and notch in posterior margin of the telson (Puls feeding frequency (Donahue 2004). 2001). Recently molted megalopae are thig- Predators: motactic, settlement is gregarious and indi- Behavior: viduals remain in high-density aggregations Bibliography into adulthood (Jensen 1989, 1991; Do- nahue 2004). Larval settlement was not ef- 1. BARNARD, L. J., D. E. BOWERS, AND E. fected by upwelling conditions, and instead C. HADERLIE. 1980. Macrura and larval abundance increased prior to spring Anomura, p. 577-593. In: Intertidal inverte- tides, suggesting tidal transport shoreward brates of California. R. H. Morris, D. P. Ab- for settlement (Mace and Morgan 2006). bott, and E. C. Haderlie (eds.). Stanford Petrolisthes cinctipes larvae do not vertically University Press, Stanford, CA. migrate and maintain their position in near- 2. CEBALLOS-OSUNA, L., H. A. CARTER, shore habitats by remaining at depth, where N. A. MILLER, AND J. H. STILLMAN. water flow would not push them offshore 2013. Effects of ocean acidification on ear- (Shanks 2009; Miller and Morgan 2013). ly life-history stages of the intertidal porce- Juvenile: Following settlement, megalopae lain crab Petrolisthes cinctipes. Journal of lose the ability to swim as their pleopods de- Experimental Biology. 216:1405-1411. generate and their body color changes (Fig. 3. DONAHUE, M. J. 2004. Size-dependent 6, Jensen 1991). competition in a gregarious porcelain crab Longevity: Petrolisthes cinctipes (Anomura, Porcel- Growth Rate: Growth occurs in conjunction lanidae). Marine Ecology Progress Series. with molting. In pre-molting periods the epi- 267:219-231. dermis separates from the old cuticle and a 4. GONOR, J. J., AND S. L. GONOR. 1973a. dramatic increase in epidermal cell growth Variations in appendage setal counts in occurs. Post-molt individuals will have soft zoea larvae of four porcellanid crabs shells until a thin membranous layer is de- (Decapoda, Anomura) from Oregon, USA. posited and the cuticle gradually hard- Crustaceana (Leiden). 25:245-252. ens. During a molt decapods have the abil- 5. GONOR, S. L., AND J. J. GONOR. 1973b. ity to regenerate limbs that were previously Descriptions of larvae of four north Pacific autotomized (Kuris et al. 2007). Porcellanid Porcellanidae (Crustacea, Anomura). Fish- crabs readily autotomize their chelipeds, to ery Bulletin. 71:189-223. avoid , and Petrolisthes cinctipes is 6. HAIG, J. 1960. The Porcellanidae no exception (Kuris et al. 2007) and autoto- (Crustacea, Anomura) of the Eastern Pa- my tends to be more common among fe- cific. Allan Hancock Pacific Expedition. male and small individuals (Wasson and 24:1-440. Lyon 2005). For complete molt staging 7. HARTMAN, B., AND M. S. HARTMAN. scheme in P. cinctipes, see Kurup 1964b. 1976. The stimulation of filter feeding in Food: A that sifts and the porcelain crab Petrolisthes cinctipes by

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]

amino acids and sugars. Comparative 15. KURUP, N. G. 1964b. The intermolt cycle Biochemistry and Physiology. 56A:19-22. of an anomuran, Petrolisthes cinctipes 8. HARVEY, A., C. B. BOYKO, P. (Randall) (Crustacea, Decapoda). Biologi- MCLAUGHLIN, AND J. W. MARTINS. cal Bulletin. 127:97-107. 2014. Anomura, p. 284-295. In: Atlas of 16. MACE, A. J., AND S. G. MORGAN. 2006. larvae. J. W. Martin, J. Biological and physical coupling in the lee Olesen, and J. T. Høeg (eds.). Johns of a small headland: contrasting transport Hopkins University Press, Baltimore. mechanisms for crab larvae in an 9. JENSEN, G. C. 1989. Gregarious settle- upwelling region. Marine Ecology Progress ment by megalopae of the porcelain Series. 324:185-196. crabs Petrolisthes cinctipes (Randall) 17. MACGINITIE, G. E., AND N. MACGINI- and (Stimpson). TIE. 1949. Natural history of marine ani- Journal of Experimental Marine Biology mals. McGraw-Hill Book Co., New York. and Ecology. 131:223-231. 18. MILLER, S. H., AND S. G. MORGAN. 10. JENSEN, G. C. 1991. Competence, set- 2013. Interspecific differences in depth tling behavior, and post-settlement ag- preference: regulation of larval transport in gregation by porcelain crab megalopae an upwelling system. Marine Ecology Pro- (Anomura, Porcellanidae). Journal of Ex- gress Series. 476:301-306. perimental Marine Biology and Ecology. 19. PAGANINI, A. W., N. A. MILLER, AND J. 153:49-61. H. STILLMAN. 2014. Temperature and 11. JENSEN, G. C., AND D. A. ARM- acidification variability reduce physiologi- STRONG. 1991. Intertidal zonation cal performance in the intertidal zone among congeners: factors regulating dis- porcelain crab Petrolisthes cinctipes. Jour- tribution of porcelain crabs Petrolisthes nal of Experimental Biology. 217:3974- spp. (Anomura, Porcellanidae). Marine 3980. Ecology Progress Series. 73:47-60. 20. PULS, A. L. 2001. Arthropoda: Decapoda, 12. KOZLOFF, E. N. 1993. Seashore life of p. 179-250. In: Identification guide to larval the northern Pacific coast: an illustrated marine invertebrates of the Pacific North- guide to northern California, Oregon, west. A. Shanks (ed.). Oregon State Uni- Washington, and British Columbia. Uni- versity Press, Corvallis, OR. versity of Washington Press, Seattle, 21. RICKETTS, E. F., AND J. CALVIN. 1971. WA. Between Pacific tides. Stanford University 13. KURIS, A. M., P. S. SADEGHIAN, J. T. Press, Stanford, California. CARLTON, AND E. CAMPOS. 2007. De- 22. RONGES, D., J. P. WALSH, B. J. SIN- capoda, p. 632-656. In: The Light and CLAIR, AND J. H. STILLMAN. 2012. Smith manual: intertidal invertebrates Changes in extreme cold tolerance, mem- from central California to Oregon. J. T. brane composition and cardiac transcrip- Carlton (ed.). University of California tome during the first day of thermal accli- Press, Berkeley, CA. mation in the porcelain crab Petrolisthes 14. KURUP, N. G. 1964a. The incretory or- cinctipes. Journal of Experimental Biology. gans of the eye-stalk and brain of the 215:1824-1836. porcelain crab, Petrolisthes cinctipes 23. RUPPERT, E. E., R. S. FOX, AND R. D. (Randall) (Reptantia, Anomura). General BARNES. 2004. Invertebrate zoology: a and Comparative Endocrinology. 4:99- functional evolutionary approach. Thom- 112. son Brooks/Cole, Belmont, CA.

Hiebert, T.C. 2015. Petrolisthes cinctipes. 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.

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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]