Pagurus Hirsutiusculus Class: Multicrustacea, Malacostraca, Eumalacostraca

Home , Crepidula adunca, Multicrustacea, Nucella emarginata, Paguristes ulreyi, Pagurus, Pagurus armatus

Phylum: Arthropoda, Crustacea

Pagurus hirsutiusculus Class: Multicrustacea, Malacostraca, Eumalacostraca

Order: Eucarida, Decapoda, Pleocyemata, Anomura Hairy hermit crab Family: Paguroidea, Paguridae

Taxonomy: The taxonomy in the Pagu- men. They have a large plate-like carapace roidea is complicated, especially among the dorsally, beneath which are five pairs of tho- genera Eupagurus, Bernhardus and Pagu- racic appendages (see chelipeds and pereo- rus (described in McLaughlin et al. 2010). pods) and three pairs of maxillipeds (see The International Commission of Zoological mouthparts). The abdomen is elongated, Nomenclature (Opinion 472) placed the ge- soft and coiled in Paguridae (Kuris et al. neric names Eupagurus and Bernhardus in 2007) (Fig. 1). the official index of invalid and rejected Cephalothorax: names, leaving the genera Pagurus Eyes: Eyestalks short, stout and with (Paguridae) and Dardanus (Diognidae) as pointed ocular scales (Wicksten 2011). valid (Hemming 1958). Thus, previous syn- Antenna: Antennal acicle usually ex- onyms for Pagurus hirsutiusculus include ceeds eyestalk in length. Chemoreceptors on Bernhardus hirsutiusculus (McLaughlin et al. antennule hairs (Barnard et al. 1980). 2010; Wicksten 2011). Mouthparts: The mouth of decapod crustaceans comprises six pairs of appendag- Description es including one pair of mandibles (on either Size: Carapace length 19–32 mm (Barnard side of the mouth), two pairs of maxillae and et al. 1980; Kozloff 1993). Puget Sound to three pairs of maxillipeds. The maxillae and 50 mm (Ricketts and Calvin 1971) and body maxillipeds attach posterior to the mouth and often extends past the margin of the shell extend to cover the mandibles (Ruppert et al. and cannot be retracted (Kuris et al. 2007). 2004). Color: Body color tan to black or green. An- Carapace: Shield (hard, anterior por- tennae dark green with white stripes. Propo- tion) wider than long (McLaughlin 1972) (Fig. dus of walking legs hairy and tipped with 1). white or pale blue and dactyls with vertical Rostrum: Triangular (Fig. 1), acute red stripes and blue spots at base. Tips of and not much longer than lateral projections chela tan or orange and walking legs have of carapace (Wicksten 2011). white band on propodus and sometimes a Teeth: Sharp medial frontal tooth blue dot. Dactyls whitish and striped with (Barnard et al. 1980). blue and red, antennal flagellum banded Pereopods: Two pairs of hairy walking with translucent and brown (Wicksten 2011) legs with dactyls about as long as propodi, or greenish with yellow spots (Barnard et al. which are banded with white. Dactyls slender 1980). Most recognizable patterns are white and about as long as propodi (Wicksten spots on antennae and white bands around 2011). Two pairs of small posterior legs are base of second and third legs (see Plate 20 adapted for holding shell. and Fig 117, Kozloff 1993) (Fig. 1). Chelipeds: Left cheliped with small General Morphology: The body of decapod hand and granular surface, slightly hairy and crustaceans can be divided into the cepha- wider than deep (Fig. 2). Right cheliped with lothorax (fused head and thorax) and abdo-

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. Pagurus hirsutiusculus. 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.

large hand, rounded, twice as wide as small Possible Misidentifications hand, granular, slightly hairy and with one Hermit crabs (superfamilies, Coenobi- large tubercule on ventral surface (Fig. 2). toidea and Paguroidea) are easily recogniza- Cheliped stout and shorter than walking ble by their unique morphology and the gas- legs, is elongated and fingers gaping in tropod shells they inhabit (although they also male. Merus and carpus with setae, gra- inhabit tubes, twigs or even bones). They use nules, spines and ridges. their last preened pears to grip the shell and Abdomen (Pleon): Abdomen asymmetrical, their soft abdomen with reduced pleopods elongate, twisted, soft and not externally and small telson and uropods. Their carapace segmented (Fig. 1). Bears small, unpaired is usually rather thin, their eyes stalked and pleopods. have pigmented corneae. They have active Telson & Uropods: Telson and uropods antennae, equipped with sensory setae. small. Telson with slightly asymmetrical Their third maxillipeds are leg-like and bear lobes and a shallow clefts laterally. Posteri- setae and they have chelae that can be large or margin with notch and spines. Uropods enough to block their external shell aperture also asymmetrical (Wicksten 2011). or sexually dimorphic. Sexual Dimorphism: Males usually larger Three hermit-crab families are currently than females (MacGinitie and MacGinitie recognized and occur on the west coast of 1949). North America: Diogenidae, Parapaguridae Shell: Usually inhabits Nassarius fossatus, and Paguridae (Wicksten 2011). Parapaguri- Nucella lamellosa (e.g. this specimen) (in dae species occur on the continental shelf. bays, Schmitt 1921), Nucella emarginata or Diogenidae species, called “left-handed” (left Littorina sp. (Kozloff 1993). Individuals often cheliped is equal to or larger than right) hermit inhabit shells of Nucella spp. except in San crabs, are generally subtidal and often inhabit Francisco Bay, where it uses shells of the shells of moon snails. The family Diogeni- gastropod species introduced from the dae includes three local species, Isocheles Atlantic. Moves to larger shells with pilosus, Paguristes ulreyi, and P. bakeri). The increased growth. Innate selection of shell Paguridae, or “right-handed” (right cheliped is is dependent on size, weight and shell larger than left) hermit crabs, on the other volume (Reese 1962) and even the potential hand, are a speciose family locally, with 10 camouflaging properties of the shell species, inhabiting the intertidal zone to the (Partridge 1980). Individuals carefully continental shelf. Nine of the 10 local pagurid examine and select appropriate shells with species belong to the genus Pagurus (Kuris et their setaceous minor chela. These setae al. 2007). The hermit crabs of the genus Pa- have sensory structures and gurus are hard to tell apart. However, many chemoreceptors that contribute to shell local species can be easily differentiated by selection (Mesce 1993). Furthermore, shell their bright red or orange antennae and, type (i.e. snail species) may be species- likewise, many are similar to P. hirsutiuseulus specific and vary throughout ontogeny in their lack of red antennae. (Straughan and Gosselin 2014). Thus, Pagurus hirsutiusculus can be distin- available shells may be a limiting resource guished from other Pagurus species in that for hermit crabs (Vance 1972; Worcester the carapace shield is conspicuously wider and Gaines 1997), but this may only be the than long and has antennae that are banded case for a specific, preferred, shell type (i.e. green and white. Furthermore, individuals are species). often not able to fully retract into their shells. 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]

They are found inhabiting the shells of Nu- local hermit crab species in the mid-littoral cella spp. and may have Crepidula spp. li- zone and within tide pools. This species has ving on the inside or outside of their shell. bright red antennae and commonly inhabits This species co-occurs with P. samuelis, but the shells of Tegula spp. Pagurus armatus is is the more common species in protected a low intertidal species, found at depths up to areas and bays. 146 m, and usually inhabits the shells of Pagurus beringanus is also a low in- Polinices spp., and are often covered with the tertidal species, found at depths up to 364 pink hydroid Hydractinia sp. Pagurus m. This species has translucent antennae caurinus is a subtidal species, to 126 m, and with a conspicuous lateral red mark and while it has a northern distribution it is rare at usually inhabits shells of Nucella lamellosa, that extent of its range. Pagurus ochetensis Ceratostoma foliata and Fusitriton is a low intertidal and subtidal species which oregonensis. It is found on rocky substrates often inhabits moon snail shells (Kuris et al. as well as sublittorally and has a whitish 2007). Their chelipeds have a red stripe, and body, red banded walking legs, and has their corneae are yellowish green. inverted V-shaped tubercules on its hands Ecological Information (Wicksten 2011; Kuris et al. 2007). Range: Type locality is Puget Sound, Wash- Pagurus quaylei is a subtidal species ington. Known range includes the Pribilof Is- (to 97 m) with antennae irregularly banded lands and Bering Strait to northern Japan and dark brown, reddish brown eyestalks (Barnard et al. 1980). Pacific Northwest to and corneae with two bands. A common Monterey, California (McLaughlin 1972). species amongst sand and polychaete Northern and southern populations used to be tubes. split into two subspecies: P. hirsutiusculus hir- Pagurus samuelis, P. hemphilli, P. sutiusculus (northern) and P. hirsutiusculus granosimanus, P. armatus, and P. caurinus venturensis (Monterey Bay, California south- all have red or orange antennae, where P. ward) (Barnard et al. 1980). These subspe- hirsutiuseulus does not. P. samuelis is a cies were split into two formal species and, high intertidal species that is common and currently, P. hirsutiusculus is replaced in its abundant on the open coast, with red southern distribution by P. venturensis antennae and carapace with white stripes. It (Wicksten 2011). inhabits shells of Tegula spp and may have Local Distribution: Coos Bay sites include Crepidula spp. living on top of or inside the South Slough, near the channel at Collver shell (Wicksten 2011). This species co- Point and the mudflat of Metcalf Preserve. occurs with P. hirsutiusculus and is Habitat: Protected areas with silt or in bays or dominant to them in terms of shell harbors (Kozloff 1993; Wicksten 2011). Tide- competition and exchange (Kuris et al. pools, under rocks (with coarse gravel), under 2007). Pagurus hemphilli is a low intertidal seaweed (Kozloff 1993). South Slough spec- and mostly subtidal species (to 50 m) with imens occur within Zostera bed in mudflats. red flagellum of antennae and corneae with Individuals appear to prefer algal cover distinct yellow rings. This species usually (Orians and King 1964) and sandy tidepools inhabits the shells of Tegula and Astraea (Reese 1962). Also present on the rocky spp, which are often themselves covered coast, in tide pools, bays and with coarse with red algae or small mollusks (e.g. sand and gravel (Kuris et al. 2007). Crepidula adunca, Acmaea mitra). Pagurus Salinity: Collected at 30, but tolerates brack- granosimanus is one of the most common

ish conditions (Barnard et al. 1980) useulus has been described (Lough 1975; Temperature: Fitch and Lindgren 1979) and proceeds via Tidal Level: Upper and middle intertidal four zoea and, a final, megalopa stage, each zone to 110 m (McLaughlin 1972; Kuris et marked by a molt (Puls 2001). Pagurus hirsu- al. 2007; Wicksten 2011). In South Slough tiuseulus zoea are shrimp-like (see paguroid at +0.15 m and -4.5 m. zoeae Fig. 53.2–3, Harvey et al. 2014: Fig. 1, Associates: In eelgrass, associates include Fitch and Lindgren; Puls 2001), with telson Littorina spp. and amphipods (South posterior having seven 7 + 7 spines, with the Slough). Associates includes those found fifth spine longest and secondary setae on the within the shell and living with the hermit inner uropod margin. Zoeal size at each crab (e.g. polynoid worms, Halosydna spp.) stage proceeds as follows (13˚C, Fitch and or sessile organisms found on top of or Lindgren 1979; McLaughlin et al. 1988): 1.9– within the shell (e.g. barnacles, limpets and 2.4 mm (Zoea I), 2.6–2.9 mm (Zoea II), 3.2– slipper shells, Crepidula spp.) (Wicksten 3.8 mm (Zoea III) and 4.1–5.2 mm (Zoea IV). 2011). Polydorid worms can infect hermit The zoea of the Paguridae are morphological- crabs heavily (Polydora commensalis). The ly similar and easiest to identify by the color parasitic isopod, Pseudione giardi, is found and distribution of their chromatophores with Puget Sound specimens (Barnard et al. (visible only in live specimens) (see Fig. 21, 1980). Other parasites and their associates Puls 2001). The megalopae have small and include the rhizocephalan parasite, reduced telson and uropods (as in adults), Peltogaster puguri (22% females infected, and chelipeds that are smooth, with no teeth 11.6% males, Alaska) as well as or hairs (see Fig. 53.7 Harvey et al. 2014. Peltogasterella gracilis and the Megalopae are 1.32 mm in length and 1.0 mm hyperparasite bopyrid isopod, Liriopsis in width and are often infested with bopyrid pygmaea in southeastern Alaska (see Fig. 1, isopod Pseudione giardi (Nyblade 1987; Puls Warrenchuk and Shirley 2000). 2001). Among competent larvae, settlement Abundance: Usually abundant in tidepools can be delayed due to lack of shells or una- (Kozloff 1993) and is one of the common vailability of food (Harvey and Colasurdo hermit crabs (MacGinitie and MacGinitie 1993; Worcester and Gaines 1997). 1949; Kuris et al. 2007). Juvenile: Antennae dark green with white stripes and walking legs white-striped, but Life-History Information never blue. Merus of both chelipeds is dark Reproduction: Male deposits sperm near brown, other leg segments are light brown the female abdomen after molting. The (Bollay 1964). sperm is stored and the female fertilizes Longevity: eggs once they are laid. Females are Growth Rate: Growth occurs in conjunction ovigerous from December through April with molting. In pre-molting periods the epi- (California, Barnard et al. 1980). Brooding dermis separates from the old cuticle and a begins in late fall and larvae hatch in dramatic increase in epidermal cell growth oc- February, with most females carrying curs. Post-molt individuals will have soft several (~ five) broods a year through spring shells until a thin membranous layer is depos- and summer months. Each brood contains ited and the cuticle gradually hardens. During up to 660 eggs and is dependent on female a molt decapods have the ability to regener- size (Fitch and Lindgren 1979). ate limbs that were previously autotomized Larva: Larval development in P. hirsuti- (Kuris et al. 2007).

Food: A detritivore, eats detritus and 6. HARVEY, A. W., AND E. A. COLASUR- scavenges for dead plant and animal DO. 1993. Effects of shell and food availa- material (Kozloff 1993). Some estuarine bility on metamorphosis in the hermit types filter plankton with their mouthparts crabs Pagurus hirsutiusculus (Dana) and (MacGinitie and MacGinitie 1949). Pagurus granosimanus (Stimpson). Jour- Predators: Other crabs. nal of Experimental Marine Biology and Behavior: Lively and active, especially shal- Ecology. 165:237-249. low water varieties (deepwater animals are 7. HEMMING, F. 1958. Official index of re- more sluggish, MacGinitie and MacGinitie jected and invalid family-group names in 1949) and will abandon shell in quiet waters zoology. First installment: names 1-273. (Ricketts and Calvin 1971). Based on a International Trust for Zoological Nomen- study with Alaskan and southern Californian clature, London. P. hirsutiusculus, it was suggested that 8. KOZLOFF, E. N. 1993. Seashore life of evolutionary shell loss may result from large, the northern Pacific coast: an illustrated active species with northern populations guide to Northern California, Oregon, (Blackstone 1989). Washington, and British Columbia. Univer- sity of Washington Press, Seattle, WA. Bibliography 9. KURIS, A. M., P. S. SADEGHIAN, J. T. 1. BARNARD, L. J., D. E. BOWERS, AND CARLTON, AND E. CAMPOS. 2007. De- E. C. HADERLIE. 1980. Macrura and capoda, p. 632-656. In: The Light and Anomura, p. 577-593. In: Intertidal inver- Smith manual: intertidal invertebrates from tebrates of California. R. H. Morris, D. P. central California to Oregon. J. T. Carlton Abbott, and E. C. Haderlie (eds.). Stan- (ed.). University of California Press, Berke- ford University Press, Stanford, CA. ley, CA. 2. BLACKSTONE, N. W. 1989. Size, shell- 10. LOUGH, R. G. 1975. Dynamics of crab living and carcinization in geographic larvae (Anomura: Brachyura) off the cen- populations of a hermit crab, Pagurus tral Oregon coast, 1969-1971. Ph.D. Ore- hirsutiusculus. Journal of Zoology. gon State University, Corvallis, OR. 217:477-490. 11. MACGINITIE, G. E., AND N. MACGINI- 3. BOLLAY, M. 1964. Distribution and use TIE. 1949. Natural history of marine ani- of gastropod shells by the hermit crabs mals. McGraw-Hill Book Co., New York. Pagurus samuelis, Pagurus granosi- 12. MCLAUGHLIN, P. A. 1972. The Hermit manus, and Pagurus hirsutiusculus at Crabs of the genus Pagurus (Crustacea, Pacific Grove California. Veliger, supple- Decapoda, Paguridae) from northwestern ment 6:71-76. North America, with a partial revision of 4. FITCH, B. M., AND E. W. LINDGREN. the genus. Ph.D. George Washington Uni- 1979. Larval development of Pagurus versity. hirsutiusculus (Dana) reared in the labor- 13. MCLAUGHLIN, P. A., R. H. GORE, AND atory. Biological Bulletin. 156:76-92. J. A. CRAIN. 1988. Studies on the proven- 5. HARVEY, A. W., C. B. BOYKO, P. zanoi and other pagurid groups: II. A reex- MCLAUGHLIN, AND J. W. MARTINS. amination of the larval stages of Pagurus 2014. Anomura, p. 284-295. In: Atlas of hirsutiusculus hirsutiusculus (Dana) crustacean larvae. J. W. Martin, J. (Decapoda, Anomura, Paguridae) reared Olesen, and J. T. Høeg (eds.). Johns in the laboratory. Journal of Crustacean Hopkins University Press, Baltimore. Biology. 8:430-450.

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