Crangon Franciscorum Class: Malacostraca Order: Decapoda Common Gray Shrimp Section: Caridea Family: Crangonidae

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Phylum: Arthropoda, Crustacea Crangon franciscorum Class: Malacostraca Order: Decapoda Common gray shrimp Section: Caridea Family: Crangonidae

Taxonomy: Schmitt (1921) described many Mouthparts: The mouth of decapod shrimp in the genus Crago (e.g. Crago crustaceans comprises six pairs of franciscorum) and reserved the genus appendages including one pair of mandibles Crangon for the snapping shrimp (now in the genus Alpheus). In 1955–56, the (on either side of the mouth), two pairs of International Commission on Zoological maxillae and three pairs of maxillipeds. The Nomenclature formally reserved the genus maxillae and maxillipeds attach posterior to Crangon for the sand shrimps only. Recent the mouth and extend to cover the mandibles taxonomic debate revolves around potential (Ruppert et al. 2004). Third maxilliped setose subgeneric designation for C. franciscorum and with exopod in C. franciscorum and C. (C. Neocrangon franciscorum, C. alaskensis (Wicksten 2011). franciscorum franciscorum) (Christoffersen Carapace: Thin and smooth, with a 1988; Kuris and Carlton 1977; Butler 1980; single medial spine (compare to Lissocrangon Wicksten 2011). with no gastric spines). Also lateral (Schmitt 1921) (Fig. 1), hepatic, branchiostegal and Description pterygostomian spines (Wicksten 2011). Size: Average body length is 49 mm for Rostrum: Rostrum straight and males and 68 mm for females (Wicksten upturned (Crangon, Kuris and Carlton 1977). 2011). Short, flattened, rounded (Fig. 2) and Color: White, mottled with small black spots, unornamented. Rostrum length reaching or giving gray appearance. exceeding the cornea of the eye (Wicksten General Morphology: The body of decapod 2011). crustaceans can be divided into the Teeth: cephalothorax (fused head and thorax) and Pereopods: Inner spine on merus of abdomen. They have a large plate-like first pereopod, hand of subchela elongate and carapace dorsally, beneath which are five dactyl closes longitudinally (versus obliquely) pairs of thoracic appendages (see chelipeds against inner spine. Second to fifth pereopod and pereopods) and three pairs of morphology as follows: second pereopod maxillipeds (see mouthparts). The body is slender and chelate; third slender and with laterally compressed and shrimp-like in the simple dactyl; fourth and fifth larger than third Caridea. The abdomen and associated and bearing setae, also with simple dactyls appendages are outstretched and the (Wicksten 2011). abdomen usually has a sharp bend (Kuris et Chelipeds: Subchelate. Dactyl al. 2007). almost parallel to hand (Plate 319A, Kuris et Cephalothorax: al. 2007) (Fig. 3). Eyes: Eyes small, pigmented and not Abdomen (Pleon): Shrimp-like, with typical covered by carapace (Wicksten 2011). caridean bend, the second segment overlaps Antennae: Antennal scale the first (Fig. 1). Male abdomen narrow (see (scaphocerite) with spine longer than blade Sexual Dimorphism). Bears rounded pleura (Fig. 2). Stylocerite (basal, lateral spine on and fifth segment with postero-lateral spine. antennule) longer than first antennule Morphology of the sixth abdominal segment is peduncle segment (Wicksten 2011). Inner taxonomically significant—in C. franciscorum, flagellum of the first antenna is greater than the sixth segment is slender, with postero- twice as long as the outer flagellum (Kuris et ventral spine and ventral groove (Wicksten al. 2007) (Fig. 2). 2011).

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12702 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] Telson & Uropods: Telson shorter than alaskensis, C. franciscorum, C. handi, C. uropods. Telson is narrow and with two pairs nigricauda and C. nigromaculata. of lateral spines, acute apex (Wicksten 2011). Crangon nigricauda, the black tailed Sexual Dimorphism: Females often have shrimp, has an antennal blade and spine of broader and larger bodies than males, which nearly equal length and cheliped fingers that have compressed and squat bodies close almost transversely. Crangon (Wicksten 2011). See also variations in nigromaculata has a striking round marking endopod morphology (Fig. 4, Siegfried 1989). on the side of the sixth abdominal segment, its fingers also close transversely, and its Possible Misidentifications range may not extend north to Oregon. The family Crangonidae is characterized by Crangon handi, from the outer coast, has a first pereopods that are subchelate and very short, stout antennal scale, and a short second pereopods (if present) that are sixth abdominal segment (Kuris and Cartlon slender and equal in size, each with 1977). Crangon alaskensis is a small shrimp, unsegmented carpus. Other characters with a slender rostrum, and lacks C. include chela dactyls that close horizontally franciscorum's very long propodus. Crangon across the end of the propodus (“like the franciscorum can further be differentiated blade of a pocket knife” Wicksten 2011), a from C. alaskensis by the chela dactyl that is rostrum that is small and without spines and a almost parallel to the hand (Plate 319A, Kuris body that is squat and somewhat depressed et al. 2007) rather than at a 45˚ angle. (although it can be broad in ovigerous Furthermore, C. franciscorum has a spine on females) (Wicksten 2011). the postero-dorsal corner of the fifth Three crangonid genera were revised abdominal segment and an inner flagellum of in 1977 by Kuris and Carlton: Lissocrangon, the first antenna that is more than two times Crangon and Neocrangon. Members of the as long as the outer (Plate 316A, Kuris et al. two former genera are found locally. These 2007). Butler calls this species Crangon genera can be differentiated by carapace franciscorum franciscorum, to distinguish it spination: Lissocrangon lacks gastric spines; from C.f. angustimana Rathbun 1902, the Crangon has one median gastric spine; and long-clawed Crangon (Butler 1980). This Neocrangon (Mesocrangon, M. munitella, latter species lives in deeper water, and within locally) has two gastric spines. a narrower range of temperatures than does Besides lacking gastric spines, C. f. franciscorum (Butler 1980). Lissocrangon species have a long, narrow and sharply recurved rostrum, a telson that is Ecological Information shorter than the uropods and a sixth Range: Type locality is San Francisco Bay, abdominal segment without a pair of dorsal California. Known range includes carinae (keel-like ridges). This genus is Resurrection Bay, Alaska to San Diego, monotypic, the only known member is L. California (Schmitt 1921; Wicksten 2011). stylirostris (Kuris and Carlton 1977; Kuris et Local Distribution: Yaquina Bay; South al. 2007). Slough (Collver Point, channel) (Krygier and Crangon species have a straight and Horton 1975). upturned rostrum and a telson that is of equal Habitat: Shallow water in bays and estuaries length or longer than uropods. The genus is with soft substrate. In bay channel with divided into groups (i.e. subgenera, Kuris and muddy and rocky substrate, also offshore in Carlton 1977) based on characters of the deeper water (Kuris et al. 2007; Wicksten sixth abdominal segment. In the first group, 2011). the sixth abdominal segment is smooth and Salinity: Euryhaline and collected at 30, lacks dorsal carinae (instead of two distal with salinity range from 0.2–34.4 (Krygier and carinae). This group is further split based on Horton 1975; Wahle 1985; Wicksten 2011; a sulcate or convex sixth abdominal segment Campos et al. 2012). ventrum. The so-called “smooth, sulcate Temperature: Great toleration of species group” (Kuris and Carlton 1977) temperature variation (e.g. 5–25˚C, Campos includes all local Crangon species, C.

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. et al. 2012) prefers warmer water than C. pleopod morphology) (Gavio et al. 1994; nigricauda (Krygier and Horton 1975). Gavio et al. 2006). Tidal Level: Intertidal to 91 m (Schimitt 1921; Larva: In caridean shrimp species Wicksten 2011). development proceeds through several zoea Associates: Collected in a trawl with Cancer larval stages (Konishi 2000; Guerao and jordani, Hermissenda sp., Rostanga pulchra, Cuesta 2014). Crangonid zoea are and other sponges. Can be infested with characterized by a wide rostrum, Bopyrid isopod Argeia pugettensis (Butler hemispherical eyestalks, antennule bases 1980). This isopod is found next to the that touch, unsegmented antennule scales branchial chamber in many crangonids and with inner flagellum having a setose spine, an forms a conspicuous bulge in the carapace exopod present on pereopod one (not 3–5), (see Plate 19, Kozloff 1993; Wicksten 2011). which is subchelate and a telson that widens The crangonid shrimps L. stylirostris and posteriorly (see Fig. 48.1, Guerao and Cuesta Crangon franciscorum may also be infected 2014; Puls 2001). Larval stages occur in the with the microspordium parasite Pleistophora plankton and earliest post-larval shrimp found crangoni, which occurs within their skeletal in brackish water of shallow tidal flats— musculature (Breed and Olson 1977). mature animals move into deeper water Crangon franciscorum individuals can also be (Israel 1936). infected with chitinoclastic bacteria that Juvenile: causes bacterial shell disease (Vibrio spp. Longevity: Females live a maximum 1 1/2 and Pseudomonas sp.). This type of bacterial years and males up to one year (Krygier and infection causes the exoskeleton to be Horton 1975; Gavio et al. 2006). darkened, cracked and amorphous, with Growth Rate: Growth occurs in conjunction underlying tissue lesions (Arnold and with molting. In pre-molting periods the Hendrickson 1997). epidermis separates from the old cuticle and Abundance: Common to abundant (Kuris et a dramatic increase in epidermal cell growth al. 2007). occurs. Post-molt individuals will have soft shells until a thin membranous layer is Life-History Information deposited and the cuticle gradually Reproduction: Many crangonid shrimp hardens. During a molt decapods have the species are considered gonochoristic, but ability to regenerate limbs that were evidence suggests that some species exhibit previously autotomized (Kuris et al. 2007). protandry (Bauer 2004). Identifying ovigerous Differential growth rate in C. franciscorum is females is easy as female crangonid shrimp large between females and males and carry eggs, which are attached with between juvenile growth ranged from 0.76–1.37 mm the joints and rami of the inner pleopods, per week (Oregon, Krygier and Horton 1975). under the abdomen and spawning has been Food: The diets of co-occurring crangonid observed almost year-round (Siegfried 1989). species, Crangon franciscorum and C. Ovigerous females collected from Dec–May, nigricauda, have been described (Wahle June or August and hatching reported from 1985). Individuals feed on small invertebrates early spring to summer (San Francisco Bay, (e.g. Neomysis mercedis, Sitts and Knight California, Israel 1936; Yaquina Bay, Oregon, 1979; amphipods, bivalves, Wahle 1985) and, Krygier and Horton 1975; Strathmann 1987; in the Columbia River estuary C. franciscorum Campos et al. 2012) during periods or in accounted for a high proportion of the total locations of high salinity (Israel 1936). A suspended particle consumption (Simenstad spawning migration was described by Krygier et al. 1990). and Horton where ovigerous females moved Predators: Demersal fishes, Dungeness to more haline and deeper water prior to crabs and harbor seals (Wicksten 2011). spawning (1975). Crangon franciscorum Commercially harvested in San Francisco exhibits protandric hermaphroditism where Bay, California for bait with L. stylirostris, C. males and females can switch from one sex nigricauda and C. nigromaculata (Israel to another and associated morphological 1936). Annual catches ranged from 320,000– changes can be observed (e.g. shape of

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12702 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] 1,360,000 kg (1940–1957) to 2,300–25,000 Crangon franciscorum (Decapoda, kg (1980s) per year (Chace and Abbott 1980). Caridea). Journal of Crustacean Behavior: Benthic shrimp that dig in the Biology. 26:295-307. sand and remain hidden with only their eyes 10. GUERAO, G., and J. A. CUESTA. and antennae visible (Wicksten 2011). 2014. Caridea, p. 250-255. In: Atlas of crustacean larvae. J. W. Margtin, J. Olesen, and J. T. Høeg (eds.). Johns Bibliography Hopkins University Press, Baltimore. 11. ISRAEL, H. R. 1936. A contribution 1. ARNOLD, J. D., and G. L. toward the life histories of two HENDRICKSON. 1997. Bacterial shell California shrimps, Crago disease in crangonid shrimp. franciscorum and Crago nigricauda. California Fish and Game. 83:118- California Department of Fish and 127. Game Bulletin. 46:1-28. 2. BAUER, R. T. 2004. Remarkable 12. KONISHI, K., and J. N. KIM. 2000. shrimps: adaptations and natural The first zoeal stage of sand shrimp history of the carideans. Animal Crangon amurensis (Brashnikov, Natural History Series. 7:1-282. 1907), with a discussion of the larval 3. BREED, G. M., and R. E. OLSON. characters of the Crangonidae 1977. Biology of microsporidan (Crustacea, Decapoda, Caridea). parasite Pleistophora crangonin n.sp. Bulletin of National Research Institute in three species of crangonid sand of Aquaculture:1-12. shrimps. Journal of Invertebrate 13. KOZLOFF, E. N. 1993. Seashore life Pathology. 30:387-405. of the northern Pacific coast: an 4. BUTLER, T. H. 1980. Shrimps of the illustrated guide to northern California, Pacific Ccoast of Canada. Canadian Oregon, Washington, and British Bulletin of Fish and Aquatic Sciences. Columbia. University of Washington 202:1-280. Press, Seattle. 5. CAMPOS, J., C. MOREIRA, F. 14. KRYGIER, E. E., and H. F. HORTON. FREITAS, and H. W. VAN DER 1975. Distribution, reproduction, and VEER. 2012. Short review of the eco- growth of Crangon nigricuda and geography of Crangon. Journal of Crangon franciscorum in Yaquina Bay, Crustacean Biology. 32:159-169. Oregon. Northwest Sciences. 49:216- 6. CHACE, F. A., D. P. ABBOTT, R. H. 240. MORRIS, and E. C. HADERLIE. 1980. 15. KURIS, A. M., and J. T. CARLTON. Caridea: the shrimps. In: Intertidal 1977. Description of a new species, invertebrates of California. Stanford Crangon handi, and new genus, University Press, Stanford, CA. Lissocrangon, of crangonid shimps 7. CHRISTOFFERSEN, M. L. 1988. (Crustacea, Caridea) from California Genealogy and phylogenetic coast, with notes on adaptation in classification of the world Crangonidae body shape and coloration. Biological (Crustacea, Caridea), with a new Bulletin. 153:540-559. species and new records for the south 16. KURIS, A. M., P. S. SADEGHIAN, J. western Atlantic. Revista Nordestina T. CARLTON, and E. CAMPOS. 2007. de Biologia. 6:43-59. Decapoda, p. 632-656. In: The Light 8. GAVIO, M. A., J. M. ORENSANZ, and and Smith manual: intertidal D. ARMSTRONG. 1994. Protandric invertebrates from central California to hermaphroditism in the bay shrimp Oregon. J. T. Carlton (ed.). University Crangon franciscorum (Decapoda, of California Press, Berkeley, CA. Caridea). Journal of Shellfish 17. PULS, A. L. 2001. Arthropoda: Research. 13:292-292. Decapoda, p. 179-250. In: 9. —. 2006. Evaluation of alternative life Identification guide to larval marine history hypotheses for the sand shrimp invertebrates of the Pacific Northwest.