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Phylum: Arthropoda, Crustacea

Crangon franciscorum Class: Multicrustacea, , Eumalacostraca

Order: , , , Common gray Family: ,

Taxonomy: Schmitt (1921) described many duncle segment (Wicksten 2011). Inner fla- shrimp in the Crago (e.g. Crago fran- gellum of the first antenna is greater than ciscorum) and reserved the genus twice as long as the outer flagellum (Kuris et for the snapping shrimp (now in the genus al. 2007) (Fig. 2). Alpheus). In 1955–56, the International Mouthparts: The mouth of decapod Commission on Zoological Nomenclature comprises six pairs of appendag- formally reserved the genus Crangon for the es including one pair of mandibles (on either sand only. Recent taxonomic de- side of the mouth), two pairs of maxillae and bate revolves around potential subgeneric three pairs of maxillipeds. The maxillae and designation for C. franciscorum (C. Neocran- maxillipeds attach posterior to the mouth and gon franciscorum, C. franciscorum francis- extend to cover the mandibles (Ruppert et al. corum) (Christoffersen 1988; Kuris and Carl- 2004). Third maxilliped setose and with exo- ton 1977; Butler 1980; Wicksten 2011). pod in C. franciscorum and C. alaskensis (Wicksten 2011). Description Carapace: Thin and smooth, with a Size: Average body length is 49 mm for single medial spine (compare to Lissocrangon males and 68 mm for females (Wicksten with no gastric spines). Also lateral (Schmitt 2011). 1921) (Fig. 1), hepatic, branchiostegal and Color: White, mottled with small black spots, pterygostomian spines (Wicksten 2011). giving gray appearance. Rostrum: Rostrum straight and up- General Morphology: The body of decapod turned (Crangon, Kuris and Carlton 1977). crustaceans can be divided into the cepha- Short, flattened, rounded (Fig. 2) and unorna- lothorax (fused head and thorax) and abdo- mented. Rostrum length reaching or exceed- men. They have a large plate-like carapace ing the cornea of the eye (Wicksten 2011). dorsally, beneath which are five pairs of tho- Teeth: racic appendages (see chelipeds and pere- Pereopods: Inner spine on merus of opods) and three pairs of maxillipeds (see first pereopod, hand of subchela elongate and mouthparts). The body is laterally com- dactyl closes longitudinally (versus obliquely) pressed and shrimp-like in the Caridea. The against inner spine. Second to fifth pereopod abdomen and associated appendages are morphology as follows: second pereopod outstretched and the abdomen usually has a slender and chelate; third slender and with sharp bend (Kuris et al. 2007). simple dactyl; fourth and fifth larger than third Cephalothorax: and bearing setae, also with simple dactyls Eyes: Eyes small, pigmented and not (Wicksten 2011). covered by carapace (Wicksten 2011). Chelipeds: Subchelate. Dactyl almost Antenna: Antennal scale parallel to hand (Plate 319A, Kuris et al. 2007) (scaphocerite) with spine longer than blade (Fig. 3). (Fig. 2). Stylocerite (basal, lateral spine on Abdomen (Pleon): Shrimp-like, with typical antennule) longer than first antennule pe-

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. . 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.

caridean bend, the second segment dominal segment without a pair of dorsal cari- overlaps the first (Fig. 1). Male abdomen nae (keel-like ridges). This genus is monotyp- narrow (see Sexual Dimorphism). Bears ic, the only known member is L. stylirostris rounded pleura and fifth segment with (Kuris and Carlton 1977; Kuris et al. 2007). postero-lateral spine. Morphology of the Crangon species have a straight and sixth abdominal segment is taxonomically upturned rostrum and a telson that is of equal significant—in C. franciscorum, the sixth length or longer than uropods. The genus is segment is slender, with postero-ventral divided into groups (i.e. subgenera, Kuris and spine and ventral groove (Wicksten 2011). Carlton 1977) based on characters of the Telson & Uropods: Telson shorter than sixth abdominal segment. In the first group, uropods. Telson is narrow and with two the sixth abdominal segment is smooth and pairs of lateral spines, acute apex (Wicksten lacks dorsal carinae (instead of two distal cari- 2011). nae). This group is further split based on a Sexual Dimorphism: Females often have sulcate or convex sixth abdominal segment broader and larger bodies than males, which ventrum. The so-called “smooth, sulcate spe- have compressed and squat bodies cies group” (Kuris and Carlton 1977) includes (Wicksten 2011). See also variations in en- all local Crangon species, C. alaskensis, C. dopod morphology (Fig. 4, Siegfried 1989). franciscorum, C. handi, C. nigricauda and C. nigromaculata. Possible Misidentifications Crangon nigricauda, the black tailed The family Crangonidae is character- shrimp, has an antennal blade and spine of ized by first pereopods that are subchelate nearly equal length and cheliped fingers that and second pereopods (if present) that are close almost transversely. Crangon slender and equal in size, each with unseg- nigromaculata has a striking round marking mented carpus. Other characters include on the side of the sixth abdominal segment, chela dactyls that close horizontally across its fingers also close transversely, and its the end of the propodus (“like the blade of a range may not extend north to Oregon. pocket knife” Wicksten 2011), a rostrum that Crangon handi, from the outer coast, has a is small and without spines and a body that very short, stout antennal scale, and a short is squat and somewhat depressed (although sixth abdominal segment (Kuris and Cartlon it can be broad in ovigerous females) 1977). Crangon alaskensis is a small shrimp, (Wicksten 2011). with a slender rostrum, and lacks C. Three crangonid genera were revised franciscorum's very long propodus. Crangon in 1977 by Kuris and Carlton: Lissocrangon, franciscorum can further be differentiated Crangon and . Members of the from C. alaskensis by the chela dactyl that is two former genera are found locally. These almost parallel to the hand (Plate 319A, Kuris genera can be differentiated by carapace et al. 2007) rather than at a 45˚ angle. spination: Lissocrangon lacks gastric Furthermore, C. franciscorum has a spine on spines; Crangon has one median gastric the postero-dorsal corner of the fifth spine; and Neocrangon (Mesocrangon, M. abdominal segment and an inner flagellum of munitella, locally) has two gastric spines. the first antenna that is more than two times Besides lacking gastric spines, Lis- as long as the outer (Plate 316A, Kuris et al. socrangon species have a long, narrow and 2007). Butler calls this species Crangon sharply recurved rostrum, a telson that is franciscorum franciscorum, to distinguish it shorter than the uropods and a sixth ab- from C.f. angustimana Rathbun 1902, the

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]

long-clawed Crangon (Butler 1980). This and Pseudomonas sp.). This type of bacterial latter species lives in deeper water, and infection causes the exoskeleton to be within a narrower range of temperatures darkened, cracked and amorphous, with than does C. f. franciscorum (Butler 1980). underlying tissue lesions (Arnold and Hendrickson 1997). Ecological Information Abundance: Common to abundant (Kuris et Range: Type locality is Bay, al. 2007). . Known range includes Resurrec- tion Bay, Alaska to San Diego, California Life-History Information (Schmitt 1921; Wicksten 2011). Reproduction: Many crangonid shrimp spe- Local Distribution: Yaquina Bay; South cies are considered gonochoristic, but evi- Slough (Collver Point, channel) (Krygier and dence suggests that some species exhibit Horton 1975). protandry (Bauer 2004). Identifying ovigerous Habitat: Shallow water in bays and estuar- females is easy as female crangonid shrimp ies with soft substrate. In bay channel with carry eggs, which are attached with between muddy and rocky substrate, also offshore in the joints and rami of the inner pleopods, un- deeper water (Kuris et al. 2007; Wicksten der the abdomen and spawning has been ob- 2011). served almost year-round (Siegfried 1989). Salinity: Euryhaline and collected at 30, Ovigerous females collected from Dec–May, with salinity range from 0.2–34.4 (Krygier June or August and hatching reported from and Horton 1975; Wahle 1985; Wicksten early spring to summer (, 2011; Campos et al. 2012). California, Israel 1936; Yaquina Bay, Oregon, Temperature: Great toleration of tempera- Krygier and Horton 1975; Strathmann 1987; ture variation (e.g. 5–25˚C, Campos et al. Campos et al. 2012) during periods or in loca- 2012) prefers warmer water than C. nigri- tions of high salinity (Israel 1936). A spawn- cauda (Krygier and Horton 1975). ing migration was described by Krygier and Tidal Level: Intertidal to 91 m (Schimitt Horton where ovigerous females moved to 1921; Wicksten 2011). more haline and deeper water prior to spawn- Associates: Collected in a trawl with ing (1975). Crangon franciscorum exhibits Cancer jordani, Hermissenda sp., Rostanga protandric hermaphroditism where males and pulchra, and other sponges. Can be females can switch from one sex to another infested with Bopyrid isopod Argeia and associated morphological changes can pugettensis (Butler 1980). This isopod is be observed (e.g. shape of pleopod morphol- found next to the branchial chamber in many ogy) (Gavio et al. 1994; Gavio et al. 2006). crangonids and forms a conspicuous bulge Larva: In caridean shrimp species develop- in the carapace (see Plate 19, Kozloff 1993; ment proceeds through several zoea larval Wicksten 2011). The crangonid shrimps L. stages (Konishi 2000; Guerao and Cuesta stylirostris and Crangon franciscorum may 2014). Crangonid zoea are characterized by also be infected with the microspordium a wide rostrum, hemispherical eyestalks, an- parasite Pleistophora crangoni, which tennule bases that touch, unsegmented an- occurs within their skeletal musculature tennule scales with inner flagellum having a (Breed and Olson 1977). Crangon setose spine, an exopod present on pereopod franciscorum individuals can also be one (not 3–5), which is subchelate and a tel- infected with chitinoclastic bacteria that son that widens posteriorly (see Fig. 48.1, causes bacterial shell disease (Vibrio spp. Guerao and Cuesta 2014; Puls 2001). Larval

Hiebert, T.C. 2015. Crangon franciscorum. 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.

stages occur in the plankton and earliest Bibliography post-larval shrimp found in brackish water of 1. ARNOLD, J. D., and G. L. HENDRICK- shallow tidal flats—mature move SON. 1997. Bacterial shell disease in into deeper water (Israel 1936). crangonid shrimp. California Fish and Juvenile: Game. 83:118-127. Longevity: Females live a maximum 1 1/2 2. BAUER, R. T. 2004. Remarkable shrimps: years and males up to one year (Krygier and adaptations and natural history of the car- Horton 1975; Gavio et al. 2006). ideans. Natural History Series. 7:1- Growth Rate: Growth occurs in conjunction 282. with molting. In pre-molting periods the epi- 3. BREED, G. M., and R. E. OLSON. 1977. dermis separates from the old cuticle and a Biology of microsporidan parasite Pleis- dramatic increase in epidermal cell growth tophora crangonin n.sp. in three species of occurs. Post-molt individuals will have soft crangonid sand shrimps. Journal of Inver- shells until a thin membranous layer is de- tebrate Pathology. 30:387-405. posited and the cuticle gradually hard- 4. BUTLER, T. H. 1980. Shrimps of the Pa- ens. During a molt decapods have the abil- cific Ccoast of Canada. Canadian Bulletin ity to regenerate limbs that were previously of Fish and Aquatic Sciences. 202:1-280. autotomized (Kuris et al. 2007). Differential 5. CAMPOS, J., C. MOREIRA, F. FREITAS, growth rate in C. franciscorum is large be- and H. W. VAN DER VEER. 2012. Short tween females and males and juvenile review of the eco-geography of Crangon. growth ranged from 0.76–1.37 mm per week Journal of Biology. 32:159- (Oregon, Krygier and Horton 1975). 169. Food: The diets of co-occurring crangonid 6. CHACE, F. A., D. P. ABBOTT, R. H. species, Crangon franciscorum and C. nigri- MORRIS, and E. C. HADERLIE. 1980. cauda, have been described (Wahle 1985). Caridea: the shrimps. In: Intertidal inverte- Individuals feed on small invertebrates (e.g. brates of California. Stanford University Neomysis mercedis, Sitts and Knight 1979; Press, Stanford, CA. amphipods, bivalves, Wahle 1985) and, in 7. CHRISTOFFERSEN, M. L. 1988. Geneal- the Columbia River estuary C. franciscorum ogy and phylogenetic classification of the accounted for a high proportion of the total world Crangonidae (Crustacea, Caridea), suspended particle consumption (Simenstad with a new species and new records for et al. 1990). the south western Atlantic. Revista Predators: Demersal fishes, Dungeness Nordestina de Biologia. 6:43-59. and harbor seals (Wicksten 2011). 8. GAVIO, M. A., J. M. ORENSANZ, and D. Commercially harvested in San Francisco ARMSTRONG. 1994. Protandric hermaph- Bay, California for bait with L. stylirostris, C. roditism in the bay shrimp Crangon fran- nigricauda and C. nigromaculata (Israel ciscorum (Decapoda, Caridea). Journal of 1936). Annual catches ranged from 320,000 Shellfish Research. 13:292-292. – 1,360,000 kg (1940–1957) to 2,300– 9. —. 2006. Evaluation of alternative life his- 25,000 kg (1980s) per year (Chace and Ab- tory hypotheses for the sand shrimp Cran- bott 1980). gon franciscorum (Decapoda, Caridea). Behavior: Benthic shrimp that dig in the Journal of Crustacean Biology. 26:295- sand and remain hidden with only their eyes 307. and antennae visible (Wicksten 2011). 10. GUERAO, G., and J. A. CUESTA. 2014.

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]

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Hiebert, T.C. 2015. Crangon franciscorum. 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.