FAMILY ACANTHEPHYRIDAE Dana, 1852 Pelagic shrimp, Red prawn

Geographic Range Acanthephyrids occur world-wide in pelagic zones. At present, many of the species are thought to occur more or less world-wide or in both the Atlantic and Pacific Oceans.

Description Acanthephyrids are pelagic shrimp, often with a thin carapace and a bright orange to red color. The rostrum varies from very short to long and toothed. The carapace may bear teeth and carinae. Some species also have teeth on the dorsal midline of the abdominal somites. Eyes may be pigmented or not. The first two pairs of pereopods are chelate and shorter than the last three pairs. The maxillipeds and pereopods bear exopods.

Molecular phylogenetic analysis on nine of the ten genera in the caridean family Dana, 1852, as well as 14 other caridean families using mitochondrial 16S and nuclear 18S rRNA genes, does not support the monophyletic status of Oplophoridae. Two disparate groups of oplophorids are revealed, with different morphological characters and ecology. It is proposed that the family Oplophoridae is restricted to the three genera Oplophorus, Systellaspis and Janicella. These three genera tend to be distributed in shallower water than the other oplophorid genera, and can also be distinguished from them by certain morphological characters. They have a thicker integument, superficial cuticular photophores and larger eyes, and the molar process of their mandibles is greatly reduced or bears a deep channel. The family Acanthephyridae Bate, 1888 is resurrected for the other seven genera, which are generally distributed in deeper water and are characterised by red soft integument, no cuticular photophores, smaller eyes and well-developed molar process of the mandibles without a deep channel. (Chan et al. 2010).Three genera , Notostomus, and , are represented along the British Columbia coast.

Acanthephya Acanthephyra chacei Acanthephyra curtirostris

Hymenodora Hymenodora acanthitelsonis Hymenodora frontalis Hymenodora gracilis

Notostomus Notostomus japonicus

References Butler, T.H. 1980. Shrimps of the Pacific coast of Canada. Ottawa; Department of Fisheries and Oceans Canada. Chan, T, Lei, H. C., Li, C. P., & Chu, K. H. 2010. Phylogenetic analysis using rDNA reveals polyphyly of Oplophoridae (: ). Invertebrate Systematics 24 (2): 172–181 Wicksten, M.K. 2012. Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces. Magnolia Press, Aukland, New Zealand. ACANTHEPHRA CHACEI Krygier & Forss, 1981

Geographic Range Southern tip of Alaskan peninsula to off the coast of Oregon.

Habitat Bathypelagic, 1500-2400 m

Morphology Exoskeleton membranous. Rostrum with 5-9 dorsal teeth, usually one ventral tooth, triangular in shape, descending from gastric region and then ascending past cornea of eye. Eye pigmented. Antenna peduncle less than half length of antennal scale; stylocerite short; scale with lateral tooth exceeding blade, blade with mid-longitudinal ridge. Third maxilliped setose, with exopod. Carapace dorsally carinate, without cervical groove, with strong lateral ridge bifurcating at anterior margin; deep depression marking branchial cavity; branchiostegal tooth on short rise, not on carina. Pereopods all with exopods; P1 chelate, short, merus with 1-2 spines; P2 chelate, longer than first, merus with 1-6 spines, ischium with 0-7 spines; P3 extremely long, dactyl small and simple, with numerous spines on carpus, merus and ischium; P4 slender, with 5-17 spines on ischium; P5 shorter, dactyl simple, with numerous spines on carpus, merus and ischium. Abdomen with A3-6 dorsally carinate, each ending in spine. Telson about as long as uropod, sulcate dorsally and usually armed with 3 pairs dorsolateral spines; with terminal spine flanked by 3 pairs lateral spines

Size Male: carapace length 37 mm; female: carapace length to 36.7 mm

Coloration Scarlet.

References Krygier, E.E. & C.A. Forss, 1981. A new Acanthephyra (Crustacea, Decapoda, Caridea) from the northeast Pacific Ocean. Journal of Biology 1: 96-104. Wicksten, M.K. 2012. Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces. Magnolia Press, Aukland, New Zealand. ACANTHEPHYRA CURTIROSTRIS Wood-Mason & Alcock, 1891 Peaked Shrimp

Geographic Range Pacific Ocean: Vancouver Island to Panama and Peru, and Japan, Philippines, Hawaii; Indian Ocean: east coast of Africa; Atlantic Ocean: Caribbean region, west coast of Africa, and Madeira Collected first in Canadian waters by G. B. Reed, WSW of Cape Scott, Vancouver Island, maximum depth 2178 m, 1964.

Habitat Bathypelagic, 300-2000 m, greatest catches at 600-900 m, does not seem to migrate daily

Morphology Body moderately deep, somewhat compressed. Shell thin but firm, surface smooth. Rostrum triangular in lateral view, as long as antennular peduncle, with 6-9 small dorsal and 1-2 ventral teeth, extending posterior as carina on anterior 2/3 of carapace. Eye small and pigmented. Antenna peduncle: short; scaphocerite long, with lateral tooth exceeding blade. Third maxilliped stout, distal segment with longitudinal ridge. Carapace with prominent carina extending posterior from branchiostegal tooth, grooves extending posteriorly from orbit and antennal region, also y-shaped groove on posterior lateral region. Pereopods: P1-2 short and chelate; P3-5 with simple dactyls. Abdomen: A2-6 with strong dorsal carina; A3-5 ending in posterior dorsal spine, A3 spine the strongest. Telson with truncate tip, 8-12 lateral spines, 4 pair distal spines; outer uropod longer than inner, and as long as or shorter than telson.

Distinctions Separated from other Acanthphrid pelagic shrimps, known to occur in British Columbia waters, by markedly compressed body, especially abdomen; rostrum as triangular frontal peak, armed with 6-9 dorsal spines, and normally 1 strong ventral spine; dorsal median carinae on A2-6 produced on all but A2 as strong posterior spines.

Size Male: total length 69 mm; female: total length to 79 mm.

Coloration Crimson.

Remarks This species is relatively rare the British Columbia coast.

References original description: Wood-Mason, J.; Alcock, A. (1891). Natural history notes from H.M. Indian marine survey steamer 'Investigator', Commander R.F. Hoskyn, R.N., commanding. No. 21. Note on the results of the last season's deep-sea dredging Ann. Mag. Nat. Hist. VII, Sixth series(XXXVIII): 186-202 Butler, T.H. 1980. Shrimps of the Pacific coast of Canada. Ottawa; Department of Fisheries and Oceans Canada. Wicksten, M.K. 2012 . Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces. Magnolia Press, Aukland, New Zealand. ______

HYMENODORA ACANTHITELSONIS Wasmer, 1972

Geographic Range Off coast of Oregon. May be found to occur in BC waters, pending future deep pelagic sampling.

Habitat Pelagic, 2400-3000 m.

Morphology Exoskeleton rather firm. Rostrum about equal to eye, with 6 dorsal teeth and continuing posteriorly as carina of carapace. Eye small, lightly pigmented, with tubercle at base of cornea. Antennae: scaphocerite with tapered blade, lateral tooth exceeding blade. Carapace: anterior area not inflated; with weak cervical groove; also supra-branchial and sub- hepatic grooves and small pterygostomial tooth. Abdomen: A1 with anterolateral lobe overlapping posterior margin of carapace. Telson with 6-7 pair dorsolateral spines, lobate end with numerous small lateral spines

Size Total length about 50 mm.

References Wasmer, R.A., 1972. A new species of Hymenodora (Decapoda: Oplophoridae) from the northeastern Pacific. Crustaceana 22: 87-91 HYMENODORA FRONTALIS Rathbun, 1902 Pacific Ambereye

Geographic Range Sea of Okhotsk and Bering Sea to southern California. Type locality west of Unalaska. First captured in British Columbia waters by the Albatross, west of Moresby Island, at 2906 m, 1889 .

Habitat Pelagic, 200-2400 m, maximum abundance at 600-1300 m. Very common in its depth range.

Morphology Exoskeleton thin, not pitted. Rostrum shorter than scaphocerite, with 3-6 dorsal and no ventral teeth, tip acute. Eye without pigment or pale amber. Antennae: stylocerite shorter than eye; scaphocerite with lateral tooth longer than blade. Third maxilliped with proximal segment somewhat concave, setose; longer than first. Carapace with suborbital and antennal spines fused into lobe; moderate branchiostegal tooth merging into carina which extends nearly to posterior margin of carapace; orbital carina running obliquely posterior to v-shaped notch, then running horizontally toward posterior margin. Pereopods: P1-2 chelate, short; P3-4 with simple dactyls, spinules on merus and ischium; P5 shorter than 4, with very short dactyl. Abdomen: somites rounded, no lateral or dorsal teeth. Telson: longer than uropods, with 6-9 pairs posterolateral spines and 6 distal spines near apex; truncate distally.

Distinctions Separated from locally known Acanthphyrids of other genera by very thin but not membranous shell; small eyes, almost colorless or amber; lack of dorsal median carina and posterior spines on all abdominal somites. Characters that set apart H. frontalis from H. glacialis and other species of the genus are its relatively long rostrum extending to or beyond end of antennular peduncle; less complex sculpturing on lateral surface of carapace.

Size Male: total length 58 mm; female: total length 53 mm.

Coloration Orange-red

Remarks H. frontalis is perhaps the most common Acanthephyrid shrimp in Canadian waters. Its apparent abundance might be attributed to a poor ability to evade trawls and plankton nets. It does not migrate vertically

References original description: Rathbun, M.J., 1902a. Descriptions of new decapod from the west coast of North America. Proceedings of the United States National Museum 24: 885-905 Butler, T.H. 1980. Shrimps of the Pacific coast of Canada. Ottawa; Department of Fisheries and Oceans Canada.

HYMENODORA GLACIALIS (Buchholz, 1874) Northern Ambereye

Geographic Range Not known with certainty, because of the earlier confusion with H. gracilus. In the North Pacific Ocean, H. glacialis probably occurs from the Bering Sea to the Gulf of Panama; also in the Okhotsk Sea. It is known from the arctic region, and in the North Atlantic Ocean southward to 30°N.

Habitat Pelagic, rarely near surface in polar seas but more common at 350-1000 m; at 2000-5610 m off Oregon.

Morphology Very similar to H. frontalis. Exoskeleton membranous, finely pitted. Carapace: anterior part swollen Rostrum barely longer than eye, lower margin convex, with 2-5 dorsal and 0-1 ventral teeth. Pereopods: P1 carpus with conspicuous distal tubercle. Telson with 5-6 pairs of dorsolateral spines. See Havens and Rork (1969) for a comparison of H. glacialis and H. gracilis.

Distinctions Distinguished from other local Acanthphyrids (including H. gracilis, if encountered) by crescent- shaped dorsoventral sulcus on lateral surface of carapace; carpus of P1 somewhat flattened and slightly setose, with a prominent distal tubercle.

Size Male: total length 48 mm; female: total length 45 mm

Coloration Blood red

Synonyms Hymenodora glauca Spence Bate, 1888 Hymenodora mollicutis Spence Bate, 1888 Pasiphae glacialis Buchholz, 1874

References from synonym: Buchholz, R., 1874. Crustaceen. In: Hartlaub, G. & M. Lindeman, Die zweite deutsche Nordpolarfahrt in den Jahren 1869 und 1870, unter Fuhrung des Kapitan Karl Koldewey: 262-399, Plates 1-15. Butler, T.H. 1980. Shrimps of the Pacific coast of Canada. Ottawa; Department of Fisheries and Oceans Canada. Havens, A., and W.L. Rork. 1969. Hymenodora glacialis (Decapoda: Natantia) from the Arctic basin. Bulletin of the Southern California Academy of Sciences 68: 19-29. Wicksten, M.K. 2012 . Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces. Magnolia Press, Aukland, New Zealand.

HYMENODORA GRACILIS Smith, 1886

Geographic Range Oregon to Baja California; off Chile and sub-antarctic waters of South Pacific, Indian and Atlantic Oceans; Gulf of Mexico.Included in key because there is a good chance that specimens will eventually be collected in BC waters

Habitat Pelagic, 300-4730 m, maximal abundance at 1250-2000 m.

Morphology Similar to H. frontalis. Exoskeleton soft and membranous. Rostrum slightly longer than eye, with 4 dorsal and no ventral teeth, apex acute. Antennae: scaphocerite with blade and lateral tooth of same length. Carapace with groove running obliquely posteriorly from orbit, with hepatic and branchial branches. Telson with 3 pairs anterolateral spines.

Size Total length about 50 mm.

Coloration Bright scarlet to red.

References original description: Smith, S.I., 1886a. Report on the decapod Crustacea of the Albatross dredgings off the East coast of the United States during the summer and autumn of 1884.— Report of the Commissioner for 1885, United States Commission of Fish and Fisheries 13: 605-705, Plates 1- 20. [preprint issued in 1886, published in journal in 1887] Wicksten, M.K. 2012 . Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces. Magnolia Press, Aukland, New Zealand. NOTOSTOMUS JAPONICUS Spence Bate, 1888 Japanese Spinyridge

Geographic Range Alaska to Oregon; Hawaii; Japan. Type locality off Honshu Island, Japan. The first known capture of N. japonicus off the Pacific coast of Canada was by the Brown Bear, at 49°50'N, at 400 m below the surface, Sept. 24, 1958.

Habitat Pelagic, 450-5380 m.

Morphology Exoseleton thin. Rostrum longer than scaphocerite, slender, with 55-83 small teeth extending posteriorly on most of carapace, 10-18 ventral teeth. Eye small, cornea well developed, small tubercle on inner surface of stalk. Antennae: short; scaphocerite with broad blade, lateral tooth exceeding blade. Third maxilliped longer than P1; Carapace arched in anterior portion, with antennal and branchiostegal teeth, two lateral carinae running posteriorly from rostrum, elongate anterior lateral carina and two posterior carinae running length of carapace from orbit to posterior margin; branchial carina extending from branchiostegal spine to posterior margin; vertical carina between anterior part of branchial carina and carina posterior to eye. Pereopods: P1-2 shorter than posterior 3 pereopods, which have simple dactyls; P5 with more setose dactyl than anterior two pairs of pereopods. Abdomen: somites all with dorsomedian carina; A3-6 with posterior dorsal spine. Telson with dorsal groove, 3-4 pair dorsolateral spines, blunt tip with 5 distal spines; outer uropod longer than inner, former reaches tip of telson.

Distinctions Known by deep carapace, with 3 prominent lateral carinae; many dorsal spines, continuous from rostral tip to near posterior margin of carapace; presence of dorsal median carinae on A1-6, those on A3-6 produced as long acute spines.

Size Males: total about 151 mm; females: total about 153 mm.

Coloration Crimson; carinae darker.

References original description: Spence Bate, C., 1888. Report on the Crustacea Macrura collected by the Challenger during the years 1873-76.— Report on the Scientific Results of the Voyage of H.M.S. ”Challenger” during the years 1873-76 24: i-xc, 1-942, Plates 1-157 Butler, T.H. 1980. Shrimps of the Pacific coast of Canada. Ottawa; Department of Fisheries and Oceans Canada. Wicksten, M.K. 2012 . Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces. Magnolia Press, Aukland, New Zealand.