Caprella Drepanochir Class: Multicrustacea, Malacostraca, Eumalacostraca

Home , Aeginina, Caprella, Caprella californica, Caprellidae, Caprelloidea, Corophiida

Phylum: Arthropoda, Crustacea

Caprella drepanochir Class: Multicrustacea, Malacostraca, Eumalacostraca

Order: Peracarida, Amphipoda, Senticaudata, A skeleton shrimp, or caprellid amphipod Corophiida, Caprelldira Family: Caprelloidea, Caprellidae, Caprellinae

Taxonomy: The Caprellidae are a very dis- the telson at the animal posterior. Caprellid tinctive family of amphipods. They were pre- amphipods differ from the rest of amphipoda viously a separate amphipod suborder, but in that the abdomen is greatly reduced, espe- were recently found to be polyphyletic, aris- cially the last three abdominal segments ing at least twice from different gammarid (urosome) and associated appendages amphipod lineages (Laubitz 1993; Takeychi (uropods). Their body is also elongated ra- 1993; Watling and Carlton 2007). Current ther than laterally compressed (compare to research places them as highly modified gammarid amphipods, e.g. Eogammarus con- members of the suborder Corophiidea fervicolus) (Kozloff 1993; Watling and Carlton (Myers and Lowry 2003; Watling and Carlton 2007). 2007), a taxon divided into two infraorders Cephalon: Round cephalon with no dorsal (Caprellida, Corophiida) each with different spines or tubercles (Fig. 1) (Laubitz 1976), evolutionary feeding strategies and associat- however body spination is a highly variable ed morphology (Myers and Lowry 2003). trait among individuals (Watling and Carlton 2007). Head partially fused with the first Description pereonite (segment of pereon) and the first Size: The illustrated specimens (from Coos pair of gnathopods (Fig. 1). Pereonite one not Bay) include a 13 mm long male (Fig. 1) and more than twice as long as head in male an 8 mm long female (Fig. 2) (Measured (Laubitz 1970) and shorter in female (Laubitz from anterior (head) to posterior (abdomen), 1970) (Fig. 2). Laubitz 1970). Males collected in Japan Rostrum: Cephalon without rostrum were 13 mm (Arimoto et al. 1976; Utinomi (Laubitz 1976). 1943) while those from Alaska were 12.4 Eyes: Small (Laubitz 1976) (Fig. 1). mm in length (Laubitz 1970). Antenna 1: Less than half total body Color: White, with brown chromatophores. length (Laubitz 1970). In males, the first an- The illustrated female is darker than the tenna is approximately equal to the cephalon male specimen. combined with pereonite two (Laubitz 1970) General Morphology: The body of amphi- (Fig. 1). Articles 2–3 of peduncle are setose pod crustaceans can be divided into three while the flagellum is shorter than peduncular major regions. The cephalon (head) or articles one and two, and bears 13 articles cephalothorax includes antennules, anten- (Laubitz 1970) (Fig. 1). In the illustrated fe- nae, mandibles, maxillae and maxillipeds male, antenna one is a little longer than ce- (collectively the mouthparts). Posterior to phalon and pereonite one and the flagellum the cephalon is the pereon (thorax) with has 10 articles (Fig. 2). seven pairs of pereopods attached to pere- Antenna 2: Antenna two in the illus- onites followed by the pleon (abdomen) with trated specimens is longer than the peduncle six pairs of pleopods. The first three sets of of antenna one and has flagellum with short pleopods are generally used for swimming, setae (Laubitz 1970) (Figs. 1, 2). while the last three are simpler and surround

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. Caprella drepanochir. 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.

Mouthparts: Mandible with molar tal to poison spine and separated by cleft. (McCain 1975) and without palp (McCain There are no anterodorsal projections on pro- 1975). Left ‘lacinia mobilis' with five teeth podus in this species (Laubitz 1970) (Fig. 1). (Fig. 5) and right ‘lacinia mobilis' denticulate The dactyl is heavy, slightly curved, with inner but not five-toothed (McCain 1975) (not figu- margin slightly denticulate and not setose. red). The gnathopod is attached just posterior to Pereon: Pereon with only six segments (not middle of pereonite two (in male) and at- seven as in other amphipods) and no pereo- tached near the middle of pereonite two, but pods on pereonites three or four not at its anterior end (in female) (Laubitz (Caprellidae, McCain 1975; Laubitz 1976). 1970) (Fig. 2). The palm of propodus has a Pereonites cylindrical and longer than deep proximal grasping spine and an accessory (Laubitz 1976). Pereonites in this species spine, and a minute distal poison spine (Fig. are without dorsal spines or tubercules, but 3). Ventral spines between insertions of are covered with fine hairs (Fig. 1). Male second gnathopods are lacking in this species pereonite one is not more than twice the (Fig. 1a). Female gnathopods are much length of the head while female pereonite smaller than those of males. one is shorter than the head (Laubitz 1970) Pereopods 3 through 7: Pereopods 5 (Fig. 2). Gills on pereonites three and four –7 prehensile (for grasping) and increase in only. Round in shape and fleshy (Caprella, size posteriorly (Fig. 1). Propodus on all Mayer 1890; Watling and Carlton 2007) pereopods rather stout, with a concave inner (Figs.1, 2). Gills in male individuals are edge and a proximal tooth with a pair of more circular and females are broadly grasping spines (Fig. 1c). Female pereopods rounded (Watling and Carlton 2007). more slender than those of males (Laubitz Oostegites (marsupium) present on 1970) (Fig. 2). pereonites 3–4 in females only. The Pleon: The pleon or seventh pereonite is re- marsupium consists of two pairs of fo- duced and often unsegmented in caprellids liaceous plates called oostegites (McCain 1975). Female individuals with one (Caprellidae, Laubitz 1976) that grow from pair of lobes, but no single-articled appendag- gill bases (Fig. 2) (Arimoto et al. 1976; Wa- es above these lobes (Figs. 2, 4) (Caprella, tling and Carlton 2007). McCain 1975). Coxae: Pleonites: Gnathopod 1: Male gnathopod one Urosomites: is small and the propodus and dactyl have Epimera: serrate grasping margins (Fig. 1) while the Telson: female gnathopod is small, setose (Fig. 2). Sexual Dimorphism: Males much larger and Gnathopod 2: Male gnathopod two is more elongate than females, with a longer very large, especially the propodus, width first pereonite and an exaggerated second being less than half overall length. The gnathopod. Females when brooding have gnathopod is setose, except the dactyl and conspicuous oostegites (see pereon) and distal part of propodus (Laubitz 1970). The lack mandible palps (Watling and Carlton basis is small, with no lateral spines at the 2007). base (Figs. 1, 1b). The propodus is Possible Misidentifications tuberculate anterodistally and palm is with In contrast to the more familiar Gam- small proximal grasping-spine, large distal maroidea, the bodies of caprellid amphipods poison spine, large triangular projection dis-

are elongate and cylindrical, their pereonites anteriorly pointed body spines and short are very long and their three pairs of pereo- spines on the stout flagellum of its second pods are prehensile. Caprellids have 2–3 antennae. This species can display ''intersex" pairs of gills on the middle pereonites and features (Laubitz 1970), making males and lack the abdominal pleopods of gammarid females difficult to distinguish. Tritella amphipods. Members of three subfamilies pilimana has laterally pointed body spines and (family Caprellidae) occur locally including, its second antennal setae are long on a Caprellinae, Paracercopinae and Phtisici- slender flagellum. It is more euryhaline than nae. The caprellid family Cyamidae are par- T. laevis and is found from Alaska to asitic on cetacean mammals. They are very California (Martin 1977). Tritella tenuissima is short bodied, dorso-ventrally flattened (like a deep water species, known off shore in isopods), and have third and fourth pereon- southern California. It lacks swimming setae ites especially adapted for hanging on to on antenna two and (some believe) should be their host. transferred to the genus Triliropus (McCain Phtisicinae have three pairs of gills, 1975). not two (unlike Caprellinae). In addition, The genus Metacaprella was they have no molar surface on the mandible. characterized by a pair of appendages above The Phtisicinae have rudimentary pereo- the usual lobes on the female abdomen pods on pereonites three and four (Laubitz (McCain 1975) where Caprella spp. have only 1970). Of this family, Perotripus brevis has the one pair of lobes (Fig. 4). Caprella been reported from California (McCain 1975; anomala and C. kenneryli were formally Watling and Carlton 2007). It, as well as members of this genus (M. anomala and M. Cercops compactus (Laubitz, 1970), occurs kenneryli). Both have a small pair of sharp in Puget Sound. Caprella compactus has spines on the heads and are reported from also been reported from the outer coast of California and from Puget Sound, Washington Oregon, at Cape Arago (Laubitz 1970; Mar- (Keith 1971; McCain 1975; Martin 1977). tin 1977) and is in the only representative of The genus Caprella is characterized by the subfamily Paracercopinae locally. Cer- the presence of gills on pereonites 3–4, cops compactus does not have an elongate oostegites and mandibles without palps body as other caprellids do, its abdomen (females) (Watling and Cartlon 2007). has five segments, and pereonites five and Caprella greenleyi has been reported living on six are short and stout (Watling and Carlton hydroids and algae and on the sea star Henri- 2007). cia spp. both in Oregon and in California The subfamily Caprellinae is the most (McCain 1969, 1975; Martin 1977). Unlike speciose with 23 species in the genus most free-living caprellids, it is quite stout, and Caprella, three in Tritella and one each in has unusual antennae-- both pairs have only Deutella and Mayerella (Watling and Carlton a uni-articulate flagellum (McCain 1975). 2007). A few caprellids have a ventral spine Tritella pereopods have only one arti- between the insertions of the second gnatho- cle and their second antennae have swim- pods (C. drepanochir does not): C. californica, ming setae (Laubitz 1970; McCain 1975). C. equilibra, C. mendax, and C. pilidigitata Three species are found in Oregon: T. laevis (Laubitz 1970). Caprella californica has a is strongly stenohaline, and is found offshore long, forward directed cephalic spine (Laubitz from British Columbia, Canada to Monterey 1970). Both the propodus and basis of the Bay, California (Martin 1977). It has male gnathopod two are very long in this

species. Caprella californica has a wide lacks grasping spines on its pereopodal distribution from the western to eastern propodi (Fig. 1c). It has been reported from Pacific coasts (Martin 1977). Caprella the coasts of Japan (Arimoto et al. 1976), Ko- equilibra has no cephalic spine (McCain rea and China (Martin 1977), and from Cali- 1975) (like C. drepanochir). But unlike the fornia (McCain 1975), but not from Puget latter species, it has anterior lateral Sound (Keith 1971) or from Oregon (Laubitz projections on pereonite five, large lateral 1970). Caprella pustulata (Laubitz 1970) has spines at the base of the gnathopod two a dorsal, upward directed knob on its head. (McCain 1975) (Fig. 1b), and the ventral The head and pereon are covered with large spines between the gnathopods (Figs. 1, and small tubercles (Keith 1971). The male is 1a). Northeast Pacific range of C. equilibra setose on the second gnathopods and on includes San Juan Islands, Washington and much of the body. The antennae have some British Columbia, Canada (Martin 1977). very long setae. Caprella pustulata is re- Caprella mendax has no cephalic spine, no ported from British Columbia, Puget Sound lateral projections on pereonite five, and and from Oregon (Laubitz 1970), but not from only small lateral spines at the bases of the California (McCain 1975; Martin 1977). second gnathopods. Its dactyl is not setose Caprella scaura (Templeton, 1836), a and its distribution ranges from Vancouver cosmopolitan species newly found in North Island, Canada to San Diego, California America (Marelli 1981), is very like C. (Martin 1977). Caprella pilidigitata has no californica above, except that it lacks a ventral lateral spine near the base of gnathopod two spine between the gnathopods, and has two and its dactyl is setose. pairs of dorsal tubercles on pereonites five One group of Caprella species has at (Marelli 1981). Pereonite four in adult males is least a slight cephalic spine (and lacks smooth dorsally in Caprella simia, a species ventral spines between the second introduced to southern California from Japan gnathopods, (as above) and includes C. (Watling and Carlton 2007). natalensis, C. penantis, C. brevisrostris, C. Obvious dorsal tuberculations on the pustulata, C. simia and C. scaura. Caprella pereonites (lacking in C. drepanochir) natalensis (=C. angusta and C. uniforma, characterize the group composed of C. Watling and Carlton 2007) has a slight alaskana, C. ferrea, C. incisa, C. mutica, C. cephalic spine and small dorsal pereonite pilipalma and C. verrucosa. Caprella alaskana spines, except on pereonite one. has quite variable dorsal pereonite spines. It Gnathopod two is attached at the anterior has long first antennae, but the flagellum is end of the second pereonite in males. The shorter than the peduncle, not longer. The northeast Pacific distribution of C. natalensis male second antenna is shorter than the first includes British Columbia, Canada to Santa two articles of the first antenna. Like C. drepa- Cruz, California (Martin 1977). Caprella nochir, C. alaskana has a first pereonite not penantis is morphologically similar to C. more than twice the length of its head (Keith natalensis however pereonite five is usually 1971). It is an intertidal species, found in longer than six and seven in the latter Alaska and British Columbia, Canada (Martin species (see Laubitz 1972; Watling and 1977). Caprella ferrea has a pair of small Carlton 2007). Caprella brevirostris has only blunt spines on its head (Laubitz 1970). The a very slightly produced rostrum, not a dorsal pereonite tubercles become large cephalic spine (Arimoto et al. 1976). It spines in the posterior pereonites (Keith differs chiefly from C. drepanochir in that it 1971). The first pereonite in the male is about

as long as the head (Keith 1971). C. ferrea which is scarcely setose (Dougherty and can be similar to C. alaskana above in its Steinberg 1953). The propodus on the second juvenile and immature stages, but not as an gnathopod in C. verrucosa is shorter than the adult (Laubitz 1970). Found in Alaska, second pereonite (Keith 1971). This species British Columbia and in Puget Sound (Keith has an anteriorly directed triangular cephalic 1971; Martin 1977). Caprella incisa has projection (Keith 1971). Found in Puget small dorsal tubercles on its pereonites, the Sound (Keith 1971), California, British Colum- propodus (on second gnathopod in males) is bia, Japan (Martin 1977). Some specimens of as long as pereonite two (Keith 1971). Its C. verrucosa from protected waters have a first antennal peduncle is finely setose ventral spine between the second gnatho- (McCain 1975). It has a triangular cephalic pods, in contradiction to most keys (Marelli projection, directed anteriorly (McCain 1981). 1975), which is lacking in C. drepanochir. C. There are two other Caprella species, incisa has been reported from British which, like C. drepanochir, have no cephalic Columbia, Canada to southern California spines, no ventral spines between the (Martin 1977). Caprella mutica, an Asian gnathopods, and no dorsal pereonite projec- species, has now been reported from tions: C. gracilior, and C. laeviuscula. California (Martin 1977; Marelli 1981), and Caprella gracilior is occasionally found was found in Coos Bay with C. drepanochir intertidally, but usually inhabits deep water (authors). It has also been called C. (below 9 m, Laubitz 1970). It has a smooth acanthogaster humboldtiensis (Martin 1977). body, except for two tubercles on pereonite Caprella mutica has dorsal projections on five. The grasping spines on the slender pereonites 3–5, but not on the anterior pereopod propodus are medial (not proximal pereonites, which are setose. It has no as in C. drepanochir). The basis of the male cephalic projections. The entire second gnathopod two is much longer than the gnathopod (males) is setose in this species propodus and the dactyl is setose (Laubitz (including the dactyl). The pereopodal 1970). It has been reported from Alaska, grasping spines (on propodus) are medial, Washington, and California, but not from not proximal as in C. drepanochir. Caprella Oregon (Laubitz 1970). Caprella laeviuscula pilipalma has low tubercles dorsally, is the most common northeastern Pacific especially on its posterior segments. It has species (Laubitz 1970), and would be a small, erect, pointed, dorsally directed expected to be found intertidally in Oregon's cephalic spine (Dougherty and Steinberg estuaries. It is the species most similar to C. 1953) and its second gnathopods are drepanochir in (according to McCain 1975, attached posteriorly to the second which does not include C. drepanochir). The pereonites in the male, and anteriorly in the main difference is in the gills: they are long female (contrast C. drepanochir). The large and oval in C. laeviuscula and round in C. propodus on the male gnathopod two has no drepanochir. The male second gnathopod in poison spine or grasping spine, but does C. laeviuscula has an extremely large poison have many long colorless hairs (Dougherty spine (it is larger in C. drepanochir). The and Steinberg 1953). Caprella verrucosa female gnathopod twp in C. laeviuscula is has large, blunt tubercles on all pereonites, attached near the middle of the pereonite it is the most tuberculate of this group. (Laubitz 1970) (contrast Fig. 2). Caprella la- Unlike many of the genus, C. verrucosa and eviuscula has a wide northern Pacific distribu- C. drepanochir have an antennal peduncle tion from Japan, to Alaska, British Columbia

and south to Monterey, California (Martin 2011) and Willapa Bay, Washington (Ferraro 1977). and Cole 2007). Caprella carina, a boreal species, ap- Life-History Information parently washed ashore in Coos Bay but its Reproduction: Development in most amphi- local establishment is unknown (Jessen pods is direct, lacking a larval stage. Little is 1969; Watling and Carlton 2007). known about the reproduction and develop- Ecological Information ment in C. drepanochir. Eggs carried by fe- Range: Original description (and presumed male in marsupium (Fig. 2), until they hatch at type region) from coast of China (Laubitz 0.4–0.5 mm. 1970; Mayer 1890; Guerra-Garcia and Larva: No larval stage is observed per se, in- Takeuchi 2003). An amphi-Pacific species stead small adult-like juveniles hatch from fe- with a range extending from (Laubitz 1970) male marsupium and grow to 1 mm long. Japan, Russia, the Arctic and Alaska as far Some stay in marsupium until mother's first as Prince William Sound. Caprella molt (Japan, Kawana, in Arimoto et al. 1976; drepanochir is an introduced species to the Wolff 2014). northeast Pacific coast and was introduced Juvenile: Some Caprella juveniles cling to in ship fouling from Asia (e.g. Japan) to San their mother’s body and grow through four Francisco Bay, California (Carr et al. 2011) molts over a period of 16 days. These juve- and Oregon (Watling and Carlton 2007). niles are protected and groomed by their Local Distribution: Coos Bay sites, inclu- mother (e.g. Caprella monoceros, Aoki and ding dock-side at the Charleston small boat Kikuchi 1991) and this extended parental care basin. has been observed in a number of caprellids Habitat: Substrate determined by food (Thiel 1997). In other species juveniles do not source as caprellids can cling to almost any cling to their mother’s body, but remain near surface. They can be found on algae, her, attached to algae, where they receive sponges, etc., but do not like sandy or mud- protection from predators and other caprellids dy bottoms (McCain 1975). (e.g. Caprella decipiens, Aoki and Kikuchi Salinity: Collected at salinities of 30 (in Co- 1991). os Bay). Longevity: Temperature: Primarily an Arctic species in Growth Rate: Amphipod growth occurs in protected, cold-temperature zones (e.g. conjunction with molting where the exoskele- Prince William Sound, AK) (Laubitz 1970). ton is shed and replaced. Post-molt individu- Tidal Level: Intertidal (Laubitz 1970) and als will have soft shells as the cuticle gradual- subtidal. ly hardens (Ruppert et al. 2004). Associates: These specimens were Caprellids undergo repeated moltings collected with Obelia sp. from floating docks, as they grow and individuals of a single spe- but can also occur with the congener cies can show great variability in size depend- Caprella mutica. In Japan, they are ing upon their age (Arimoto et al. 1976). Sex- commonly associated with Tubularia sp. ually mature females are 7.5 mm in length, Abundance: Locally common in Coos Bay while males are 18 mm (Arimoto et al. 1976). (Charleston boat basin), especially in July. Food: Caprellids can eat many things by dif- One of the most abundant epifaunal species ferent methods. Presence of plumose setae in eelgrass (Zostera marina) communities in on second antennae provides the ability to San Francisco Bay, California (Carr et al. filter food and to scrape periphyton from sur-

faces to which they cling (Caine 1977) (e.g. ciety of Washington. 66:39-49. Obelia, in Coos Bay). Some individuals will 6. FERRARO, S. P., and F. A. COLE. 2007. nip off hydroid polyps as well as diatoms or Benthic macrofauna habitat associations detritus (Kozloff 1993). When feeding, the in Willapa Bay, Washington, USA. Estua- caprellid hangs on with prehensile pereo- rine Coastal and Shelf Science. 71:491- pods and uses antennae and gnathopods 507. for eating. 7. GUERRA-GARCIA, J. M., and I. Predators: Caprellids are fed upon by bot- TAKEUCHI. 2003. The Caprellidea tom fishes (cod, blennies, skates, sea bass), (Malacostraca, Amphipoda) from Mirs Bay, also by shrimp, anemones (McCain 1975), Hong Kong, with the description of a new and hydroids (e.g. Candelabrum fritchmanii, genus and two new species. Journal of Hewitt and Goddard 2001). Crustacean Biology. 23:154-168. Behavior: Movement is inchworm-like: 8. HEWITT, C. L., and J. H. R. GODDARD. grasping substrate with large anterior gnath- 2001. A New species of large and highly opods, then pulling up posterior and grab- contractile hydroid in the genus Candela- bing on with pereopods and posterior ap- brum (Hydrozoa : Anthoathecatae) from pendages. southern Oregon, USA. Canadian Journal of Zoology. 79:2280-2288. Bibliography 9. JESSEN, M. P. 1969. The ecology and 1. AOKI, M., and T. KIKUCHI. 1991. Two taxonomy of the Caprellidae (order: Am- types of maternal care for juveniles ob- phipoda; Suborder: Caprellidea) of the served in Caprella monoceros (Mayer, Coos Bay, Oregon, area. Ph.D. University 1890) and Caprella decipiens (Mayer, of Minnesota. 1890) (Amphipoda, Caprellidae). Hydro- 10. KEITH, D. E. 1971. Substrates election in biologia. 223:229-237. caprellid amphipods of Southern Califor- 2. ARIMOTO, I., D. KYOTO, and J. SETO nia, with emphasis RINKAI. 1976. Taxonomic studies of on Caprella californica Stimpson and Caprellids (Crustacea, Amphipoda, Caprella equilibra Say (Amphipoda). Pacif- Caprellidae) found in the Japanese and ic Science. 25:387-394. adjacent waters. Nihon Warekararui 11. KOZLOFF, E. N. 1993. Seashore life of Shuppanbu, Yokohama. the northern Pacific coast: an illustrated 3. CAINE, E. A. 1977. Feeding mechanisms guide to northern California, Oregon, and possible resource partitioning of the Washington, and British Columbia. Univer- Caprellidae (Crustacea: Amphipoda) sity of Washington Press, Seattle, WA. from Puget Sound, USA. Marine Biology. 12. LAUBITZ, D. R. 1970. Studies on the 42:331-336. Caprellidae, (Crustacean, Amphipoda) of 4. CARR, L. A., K. E. BOYER, and A. J. the American North Pacific. National Mu- BROOKS. 2011. Spatial patterns of epi- seum of Natural Sciences (Ottawa) Publi- faunal communities in San Francisco Bay cations in Biological Oceanography:1-89. eelgrass (Zostera marina) beds. Marine 13. —. 1972. The Caprellidae (Crustacea, Am- Ecology. 32:88-103. phipoda) of Atlantic and Arctic Canada. 5. DOUGHERTY, E. C., and J. E. STEIN- National Museum of Natural Sciences BERG. 1953. Notes on the skeleton (Ottawa) Publications in Biological Ocean- shrimps (Crustacea: Caprellidae) of Cali- ography: 1-82. fornia. Proceedings of the Biological So- 14. —. 1976. On the taxonomic status of the

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