Larval development of the grooved tanner crab, chionoecetes tanneri rathbun, 1893 (decapoda: Brachyura: Majidae) described from the laboratory‐reared specimens
Larval development of the grooved tanner crab, chionoecetes tanneri rathbun, 1893 (decapoda:...
Hong, Sung
Yun; Park, Wongyu; Perry, R. Ian; Boutillier, James A.
2009-01-01 00:00:00
Animal Cells and Systems 13: 59-69, 2009 Larval Development of the Grooved T anner Crab, Chionoecetes tanneri Rathbun, 1893 (Decapoda: Brachyura: Majidae) Described from the Laboratory- reared Specimens 1 2 2 Sung Y un Hong, W ongyu Park *, R. Ian Perry , and James A. Boutillier Department of Marine Biology , Pukyong National University , Pusan 608-737, Korea; Department of Marine Biotechnology , Soonchunhyang University , Asan 336-745, Kor ea; Department of Fisheries and Oceans, Pacific Biological Station, Nanaimo, B.C. V9T 6N7, Canada Abstra ct: This p aper document s the defining morphological INTRODUCTION characteristics of the larval stages of Chionoecetes tanneri Rathbun, 1893, the grooved Tanner crab, from specimens Chionoecetes tanneri Rathbun, 1893, the grooved Tanner reared in the laboratory . Chionoecetes tanneri larval stages crab, is a majid crab ranging in distribution from Kamtchatka, include two zoeae and one megalopa. The first zoea is Russia to Cortez Bank, Mexico (Schmitt, 1921; Garth, characterized by: six setae on the posterior margin of the 1 958; Ha r t , 1982 ) . Chionoecetes tanneri has been considere d carapace; postero-lateral spines on abdominal somites 3 and 4, extending beyond the posterior margin of adjacent for commercial exploitation in at least some parts of its somites and bearing 9-10 spinnules; 12 plumose setae and range, including the offshore waters of British Columbia, one stout distal plumose seta present on the margin of the Canada (Pereyra, 1966, 1968, 1972; Jamieson, 1990; scaphognathite of the maxilla; and one fused lateral spine Jamieson et al., 1990; Workman et al., 2002), thereby and one articulated dorso-medial spine on each fork of the providing part of the motivation for this study. Their telson. The second zoea is characterized by: 9 setae on the expected distribution is along the continental slope at postero-lateral margin of the carapace; a serrated mandible depths of 458-1784 m (Hart, 1982) and they do not appear molar; a mandibular palp bud; 25-26 plumose setae on the margin of the scaphognathite of the maxilla; pereiopods with to occur in coastal waters (Jamieson, 1990). They have well-developed gills and buds; and four pairs of stout setae been observed to occur in fine sediment habitats (Pereyra, on the posterior margin of the telson. For the megalopal 1966; Hart, 1982), and in aquaria have been observed to stage, the distinguishing characteristics include: a rostral feed on brittle stars and pieces of fish. In Oregon waters, spine equal in length to the supraorbital spine; six setae on they have been observed to release eggs, moult and mate the exopod of the uropod; and a single spine on the ischium during early spring, with reproduction taking place on an of the second pereiopod. This study allows C. t anneri larvae to be distinguished from the larvae of known sympatric annual cycle (Pereyra, 1966). However, considering their congeners. This information provides a basic taxonomic tool wide distribution range, they may have various reproductive for researchers in fisheries management and zooplankton schedules throughout the distribution range, as indicated in ecology who are addressing issues related to trophic Dungeness crab in the west coast of North America (Park et interactions, metapopulation dynamics and ecosystem al., 2007; Park and Shirley, 2008). Largely because of impacts in the evolving marine resource management identification prob le ms, larvae of g rooved T anner crab have strategies in the North Pacific, and those related to not been observed, but are assumed to occur near surface C h io noece te s species in particular. waters during late spring and summer; newly settled Key words: larval development, Chionoecetes tanneri , mega- benthic juveniles were observed below the depths of adult lopa, zoea, grooved tanner crab crabs (Pereyra, 1966). Chionoecetes contains five species: the Snow crab, C. * T o whom correspondence should be addressed. opilio; the Red queen crab, C. japonicus ; the Tanner crab, T el: +82-41-530-1282; Fax: +82-41-530-1638 E-mail address: pwg09@hotmail.com C. bairdi ; the grooved Tanner crab, C. tanneri ; and the ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 59 Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier triangle Tanner crab, C. angulatus. Of these, the larvae of formalin an d dissected in 10 0% ethylene glycol. The larvae C. opilio and C. japonicus have been described from were dissected using a M5 Wild binocular microscope. Japanese waters (Aikawa, 1937; Kurata, 1963; Kon, 1980; Appendages were illustrated using a M1 1 W ild microscope Konishi and Matsumoto, 2002), and those of C. opilio and equipped with a drawing tube. Measurements of key C. bairdi have been described from Alaskan waters morphological characteristics included: dorsal spine length (Haynes, 1973; Jewett and Haight, 1977). Larvae of C. (DSL); rostral spine length (RSL); distance between tips of tanneri were illustrated from plankton samples taken in dorsal and rostral spines (RDSL); lateral spine length Oregon waters (Lough, 1975); however, the larvae of C. (LSL); distance between tips of lateral spines (LLSL); tanneri have been difficult to distinguish from its sympatric carapace length as measured from the posterior margin of congeners because of the similarity and overlap of a the eye socket to the mid-dorsal posterior margin of the number of morphological characters. W encker et al. (1982) carapace for zoeae and from the tip of the rostrum to mid- reported finding some rare and unknown Chionoecetes dorsal posterior margin of the carapace for megalopa (CL). larvae of deep-dwelling Chionoecetes species from the A min i mu m of fi v e sp e ci me ns we re ex amin ed a n d me a sure d regions overlaying the co ntinental slope of the south eastern for each larval stage. Specimens used for the research have Bering Sea. Considering the deep-dwelling habit ranges of been deposited in the Invertebrate Laboratory, Pukyong C. tanneri and its larval characteristics, which did not National University, Pusan, Korea as PKNU-INVERT con form to th e published characteristics of C. bair di and C.number 02-01. opilio, it was postulated that the unknown Chionoecetes larvae may have been the larvae of C. tanneri or some other RE SU L T S deepwater Chionoecetes species. The objective of the present study is to describe the zoeae The larval development of C. tanneri Rathbun comprised and megalopa of C. tanneri reared in the laboratory and to two zoeal an d one megalopal stages. No meg alopa mo ulted compare them with previously described larvae of knownto the first crab stage. sympatric congeners. This study is intended to provide a taxonomic tool for researchers studying zooplankton inFirst zoea (Fig. 1) general and Chionoecetes species in particular to address questions of trophic interactions, metapopulation dynamics Size: CL=1.29 mm (1.20-1.38); DSL=2.40 mm (2.25- and ecosystem impacts in the evolving marine resource2.58); RSL=1.90mm (1.73-2.03); management strategies in the North Pacific.RDSL, 5.73mm (5.48-6.08); LSL=1.44mm (1.35-1.53). Duration: mean 30.7 (range 26-53) days. MA TERIALS AND METHODSColor: dark vermillion or dark crimson, chromatophores on abdominal somites 2-4, precardiac region and frontal st An ovigerous female C. tanneri was collected during a base of dorsal spine of the carapace, protopodite of 1 deep water assessment survey aboard the CCGS W. E.maxilliped, and mandibles. Ricker on 11 February 2000 from a crab pot fished at a Carapace (Fig. 1A, B). Dorsal, rostral, and lateral spines depth of about 400 m near Brooks Peninsula, Vancouver present; dorsal and rostral spines armed with numerous o o Island, British Columbia, Canada (128 00'W, 50 1 0 'N). In spinules on distal third; dorsal spine slightly longer than situ water temperature was around 5 C. The female was rostral spine; rostral spine slightly posteriorly curved; transported to the laboratory and maintained in a 400 L lateral spines, almost half as long as dorsal spine, armed aquarium at a constant temperature (8.8 C) until the larvae with numerous spinules, at right angle to carapace, curved hatched. slightly downward; a minute frontal knob between eyes; a Hundreds of larvae hatched from the female on 16 pair of antero-median and a pair of minute postero-dorsal February 2000. Of these, a single larva was transferred for setae; postero-lateral margin fringed with six submarginal rearing into each of one hundred 250 mL jars. An additional setae; margin with one plumose anterior seta and five 150 larvae were reared for mass culture in a 20 L jar. The posterior setae; lateral margin with two deep indentations; mass culture samples were used for drawing and descriptions. one just posterior to eye, one on the middle of the lateral Newly hatched Artemia nauplii were fed to the larvae.margin. Water temperature during the larval rearing ranged from Eyes (Fig. 1A, B). Tag eyes sessile onto the end of the o o o 8.8C through 10.4C (mean 9.6C); salinity from 26.0‰ tocarapace. 31 .0 ‰ (mean 28.9‰). W ater in each jar was changed daily Antennule (Fig. 1C). Uniramous; with 2 setae and 2 after examining the condition of the larvae, followed byaesthtascs. feeding with Artemia nauplii. Antenna (Fig. 1D). Biramous ; spinous process (protopod) Exuviae and specimens were preserved in buffered 5% elongate, subequal in length to rostral spine; armed with 60 ANIMAL CELLS AND SYSTEMS Vol. 13 No. 1 Larv al Dev elopment of the Groov ed T anner Crab numerous spinules; endopod rudimentary, a minutedistal segment. dendrical; exopod slender; less than one-third length of the Antenna (Fig. 2D). Endopod as an elongate bud, one- spinous process; two subterminal spines near acute tip. quarter the length of the spinous process (protopodite); Mandible (Fig. 1E). Incisor and molar processes well-exopod about one-half the length of spinous process. developed; palp absent. Mandible (Fig. 2E). Molar serrations, truncated and Maxillulae (Fig. 1F). Coxal endite with 2+6 setae; basial fused; both mandibles with a median indentation; endite with three fused plumodenticulate cuspidate setae, amandibular palp bud present. setal bud on the distal corner, and four setae; endopod 2- Maxillule (Fig. 2F). Basial endite with four fused segmented with 1, 2+4 plumodenticulate setae. plumodenticulate cuspidate setae and five setae; plumose Maxilla (Fig. 1G). Coxal endite, bilobed with 4 + 4 setae;exopod seta on outer margin; otherwise unchanged. basial endite, weakly bilobed with 5+5 setae; endopod, Maxilla (Fig. 2G). Basial endite with 5+6 setae; exopod bilobed with 3 + 3 setae; exopod (scaphognathite) margin(scaphognathite) margin with 25-26 plumose setae. with 12 plumose setae and one distal stout plumose seta. First maxilliped (Fig. 2H). Coxa without epipod bud; First maxilliped (Fig. 1H). Coxa with a denticulate seta; basis with an epipod bud and nine medial denticulate setae, basis with nine medial denticulate setae arranged 2, 2, 3, 2; arranged 2, 2, 3, 2; endopod setation unchanged; exopod endopod 5-segmented with 3, 2, 1, 2, 5 (1 subterminal + 4with six plumose natatory setae; otherwise unchanged. terminal) setae; exopod with four terminal plumose Second maxilliped (Fig. 2I). Basis with five medial natatory setae. denticulate setae; exopod with six plumose natatory setae; Second maxilliped (Fig. 1I). Coxa without setae; basisotherwise unchanged. with 4 medial denticulate setae arranged 1, 1, 1, 1; endopodThird maxilliped (Fig. 2J). Biramous epipod; with gill. 3-segmented with 1, 1, 5 (2 subterminal+3 terminal) setae;Pereiopods (Fig. 2K). Well-developed; with gill buds. exopod with 4 terminal plumose natatory setae. Abdomen (Fig. 2A, L). Six somites plus telson; somites Third maxilliped (Fig. 1I). Present; biramous. 2-6 with a pair of mid-dorsal setae and a pair of pleopod Pereiopods (Fig. 1I). Present; chela bilobed. buds; the posterolateral spine on somites 3-5 naked and Abdomen (Fig. 1A, J). Five somites plus telson; somite 1 barely extended beyond the posterior margin of the with (or rarely without) a pair of mid-dorsal setae; somites adjacent somites; somite 6 with a pair of short 2-3 with a pair of small, subequal, dorsolateral processes;posterolateral spine. somites 2-5 with a pair of posterodorsal setae. Somites 3-5 Telson (Fig. 2A, B, L). Telson fork with an articulated with a pair of posterolateral spines; the spines on somites 4-lateral spine and an articulate dorsomedial spine; furca with 5, armed with 9 or 10 spinules and extended beyond four pairs of stout posterior marginal setae of which the posterior margins of adjacent somites; spine on somite 5,innermost pair is much smaller than the others. n a k ed . T elson (Fig. 1A, J). T elson fork long and flexed distallyMegalopa (Figs. 3 and 4) with a fused lateral spine and an articulate dorsomedial spine; mid-posterior margin deeply concave with a medialSize: CL=3.25mm (2.90-3.55). arch; furca with three pairs of stout posterior marginal Duration: eleven megalopae obtained from 23 second setae. zoeae; survived for 45 days; no megalopas molted to first crab. Second zoea (Fig. 2) Carapace (Fig. 3A, B). Longer than wide; rostrum directed anteroventrally with a pair of stout anterolateral Size: CL=1.83 mm (1.25-2.05); DSL=2.59 mm (2.25- processes; protogastric region with a pair of stout dorsal 3.05 .); RSL = 1.96 mm (1.83-2.13); RDSL = 5 .5 1 mm (5.35- spines; branchial and metagastric regions elevated; 5.70); LSL=0.96mm (0.83-1.05).intestinal region with a pair of stout dorsal processes; lateral Duration: 30.6 (29-34) days branchial regions with a pair of lateral protuberances; six Color: Similar to first zoea.pairs of setae on protogastric and intestinal regions. Carapace (Fig. 2A, B). Lateral spine almost a quarter ofEyes (Fig. 3A). Stalked. dorsal spine; ventral margin with an indentation posterior to Antennule (Fig. 3C). Peduncle 3-segmented with 0, 2, base of antenna; posterolateral margin fringed with nine 3 + 1 setae; endopod 2-segmented with 0,4 setae; flagellum submarginal or marginal setae.3-segmented 0, 4+1, 8 aesthetascs, and setae. Eyes (Fig. 2A, B). Stalked. Antenna (Fig. 3D). Peduncle 4-segmented with 0,0,1,4 Antennule (Fig. 2C). 3-segmented; proximal segmentsetae; flagellum 5-segmented with 0,3,4,0,3 setae. with a minute seta on the basal outer margin, one aesthetasc Mandible (Fig. 3E). Cutting edges well-developed, not on the penultimate segment, and eight aesthetascs on the clearly divided into incisor and molar processes; palp 2- ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 61 Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier Fi g. 1. T he gr oov ed T anner c rab, Chonoec etes t anner i Rat hbun, 1893, fir s t z oea. A, l ateral v i ew ( ab); B, f ront al v i ew ( ab); C, ant ennul e ( ae) ; D, antenna (ac); E, mandibles (ae); F , maxillule (ae); G , maxilla (ae); H, first maxilliped (ac); I, second maxilliped (ac); J, ab domen and telson (ad). Scale bars with dif ferent sizes at the lef t bottom indicate 0.5 mm. segmented with 0, 12 setae. endite with 4 + 20 setae; endopod 2-segmented with 0, 3 Maxillule (Fig. 3F). Coxal endite with 14 setae; basialsetae. 62 ANIMAL CELLS AND SYSTEMS Vol. 13 No. 1 Larv al Dev elopment of the Groov ed T anner Crab Fi g. 2. The grooved Tanner crab, C hon oec etes t anner i Rat h bun, 18 93, s e c ond z oea. A, l a teral vi ew ( a b) ; B, fr on t a l v i ew (ab); C, a n t enn ul e (ae); D, antenna (ad); E, mandibles (ae); F , maxillule (ae); G , maxilla (ae); H, first maxilliped (ac); I, second maxi lliped (ac); J, thi rd maxi lliped (ae); K, pereiopod buds (ad); L, abdomen and telson (ac). Scale bars with dif ferent sizes at the left bottom indicate 0.5 mm. Maxilla (Fig. 3G). Coxal endite bilobed with 1 1+6 setae; process with 14 long setae; coxal and basial endites with basial endite bilobed with 9+10 setae; endopod with a numerous setae; endopod not distinctly segmented, distal naked process and 6 marginal plumose setae; scaphognathite part flattened, with two setae; exopod (scaphognathite) 2- fringed with 64-70 plumose setae.segmented with 1, 4 plumose setae. First maxilliped (Fig. 4A). Epipod, a long blade-like Second maxilliped (Fig. 4B). Coxa with one epipod and ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 63 Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier F ig. 3. T h e gr ooved T anner c r a b , Chonoecet es t a n ner i Rat h bun, 1893, megalop a. A, lat e r a l v i ew ( ab); B, dors a l v i ew (ab), C, antennule ( ad); D , a nt en na (ad); E, m andible (ae); F , maxill u l e (af ) ; G , maxilla (ae); H, later a l v i ew of abdomen (ac ) . Sc ale b a r s wi th dif f er ent s iz es at the lef t bottom ndicate 0.5 mm. one gill bud, epipod with five long setae; endopod 5- exopod 2-segmented, distal segment with six terminal segmen te d with 0,1,1,4,9 setae; exopod 2-segmen ted , distalplumose setae. segment with 0,4 plumose setae. Pereiopods (Fig. 4D-H). Pereiopods 1-3 with gills; all Third maxilliped (Fig. 4C). Coxa with 11 setae and one segmented and sparsely spinous; pereiopod 2 with a spine epipod with 30 long setae; basis/ischium with 3/25-30on basipodite. spines; merus with 10-11 spines; carpus with six spines;Abdomen (Figs. 3A, B, H). Six somites with 2, 5, 4, 4, 1, propodus with 7-8 spines; dactylus with seven spines; 1 pairs of surface setae; posterolateral mar gin of so mite 2-5 64 ANIMAL CELLS AND SYSTEMS Vol. 13 No. 1 Larv al Dev elopment of the Groov ed T anner Crab Fig. 4. The grooved T anner crab, Chonoecetes tanneri Rathbun, 1893, megalopa. A, first maxilliped (ac); B, second maxilliped (ac); C, third maxilliped (ac); D-H, pereiopod 1-5 (ab); I, pleopod on abdominal somite 1 (ac). Scale bars with different sizes at the left bottom indicate 0.5mm . broadly truncated.Telson (Fig. 3A, B, 4J): Longer than wide; naked margin. Pleopods (Figs. 3A, H, 4I). Present on abdominal somite 2-5; endopod, a simple rod-like bud with 2 or 3 curved DISCUSSION processes; exopods of pleopod 2-5 with 6,7,6,7 plumose natatory setae. Of the five Chionoecetes species in the Pacific Ocean, the Uropods (=pleopod on somite 6) (Figs. 3A, B, 4J): larvae of C. opilio, C. japonicus , and C. bairdi have been Uniramous with six plumose natatory setae. described, while those of C. tanneri and C. angulatus have ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 65 6 S 6 u T able 1. Comparison of the larval characteristics of Chionoecetes first zoeae C. opilo elongatus C. opilio C. opilio C. opilio C. japonicus C. japonicus C. bairdi C. tanneri W (Konishi and (Jewett and Haight, (Kurata, 1963) (Haynes, 1973) (Motoh, 1973) (Kon, 1980) (Motoh, 1976) (The present study) Matsumoto, 2002)1977) Carap ace Carapace length (CL) 1.8-2.0 mm 0.54 (0.45-0.73) mm 1.83 (1.25-2.02) mm Dorsal spine length (DSL) 2.3 mm 5.33 (5.04-5.78) mm 2.59 (2.25-3.05) mm Rostral spine length (RSL) 1.8-2.0 mm 1.96 (1.83-2.13) mm Dorsal-rostral spine length (DRSL) 4.5-4.9 mm 4.6-5.4 mm 4.6-5.2 mm 5.21 (4.85-5.52) mm 1.24 (1.11-1.39) mm 4.17 (3.96-4.55) mm 5.51 ( 5.35-5.70) mm Lateral spine length (LSL) 1.3 mm 0.96 (0.83-1.05) mm Lateral-Lateral spine length (LLSL) 3.3-3.6 mm 3.3-3.5 mm 3.69 (3.46-3.93) mm 2.73 (2.52-2.79) mm Posterolateral margin seta 3* 3** 5 6 6 Antennule3+22+152+13+22+2 Antenna i Exopod/Spinous process <1/2 1/4 Endopodite A minute denticle minute tubercle A small protunerance A minute denticle Maxillule Coxal endite setae 7 7 7778 Basial endite setae 6 or 7 7 7 7 7+1 7+1 Endopod setae 1,6 1,6 1,6 1,6 1,6 1,6 Maxilla Coxal endite setae 4+4 4+4 4+4 4+4 4+4 4+4 Basial endite setae 5+(4 or 5) 5+5 5 (or 6)+5 (or 4) 5+6 5+5 5+5 Endopod setae 3+3 3+3 3+3 3+3 3+3 3+3 Scaphognathite setae 15+1 (10 or 11)*(2 or 3) (11-13)+1 12 or 13 12 or 13 11 (or rarely 12)+1 12+1 First maxilliped Basial setation 2+2+3 2+2+3+3 2+2+3+3 2+2+3+3 3+3+2+2 2+2+3+2 N Endopod setation 3,2,1,2,5 3,2,1,2,5 3,2,1,2,5 3,2,1,2,5 3,2,1,2,5 2,1,1,2,5 Second maxilliped Basial setation 1+1+1+1 1+1+1+1 1+1+1+1 1+1+1+1 1+1+1+1 1+1+1+1 Endopod setation 1,1,5 1,1,5 1,1,5 1,1,5 1,1,5 1,1,5 1 A Table 1. Continued C. opilo elongatus C. opilio C. opilio C. opilio C. japonicus C. japonicus C. bairdi C. tanneri (Konishi and (Jewett and Haight, (Kurata, 1963) (Haynes, 1973) (Motoh, 1973) (Kon, 1980) (Motoh, 1976) (The present study) Matsumoto, 2002)1977) Abdomen YSomite 1-5 or somite Dorsal paired setae on somites Somite 1-5 Somite 2-5 Somite 1-5 2-5 Dorsolateral process on somite 2 Straight Straight Slightly curved Slightly curved Straight l Dorsolateral spine on somite 3 Straight Slightly curved Slightly curved Straight Barely extending Extending a little Extending beyond beyond the bit beyond the Extending beyond Extending beyond Extending beyond Extending beyond the posterior Posterolateral spine on somite 3posterior posterior the posterior the posterior margine the posterior the posterior margine margine of somite margine of margine of margine of somite 4 of somite 4 margine of somite 4 of somite 4 somite 4 somite 4 Not quite Extending beyond E x ten d in g a litt l e Extending beyond Extending beyond Extending beyond Extending beyond reaching the the posterior bi t b e yon d th e Posterolateral spine on somite 4 the posterior the posterior margine the posterior the posterior margine poterior margine margine of somite posterior margine margine of somite 5 of somite 5 margine of somite 5 of somite 5 of somite 5 5 of so mite 5 Numerous Covered by 1 or 1 spinules 7 Spinules Spinules Spinules Few spinules 9-10 spinules spinules Posterolateral spine on somite 5SmoothSmooth Telson Lateral fused spine 1 1 1 1 1 1 1 Minute secondary spine 1 1 1 1 None Dorsomedial articulated spine 1 1 1 1 1 1 1 *Motoh (1973, Fig. 1B), **Motoh (1976, Fig. 1C) Table 2. Comparison of the larval characteristics of Chionoecetes megalopae v C. opilo elongatus C. opilio C. opilio C. japonicus C. japonicus C. bairdi C. tanneri (Konishi and Matsumoto, (Jewett and Haight, (Kurata, 1963) (Motoh, 1973) (Kon, 1980) (Motoh, 1976) (The present study) p 2002)1977) m Carapace Carapace length (CL) 3.6 mm 2.9-3.3 mm 3.0 mm 2.89-3.37 mm 2.83-3.26 mm 3.12-3.48 mm 2.90-3.55 mm Equal to theLonger than the Equal to the Equal to the Equal to the Three times longer Equal to the e Rostral spine lengthsupra-orbitalsupra-orbital supra-orbital supra-orbitalsupra-orbital than the supra-orbital supra-orbital c spinespinespinespinespinespinespine A minute lateral spine on AbsentAbsentAbsentAbsentAbsentPresentAbsent pterygostomial-branchial region Spine on basipodite of preiopds 0,1,1,1,01,1,1,1,0?0,1,1,1,00,1,1,1,01,1,1,1,00,1,0,0,0 1,2,3,4,5 Uropod Exopod seate 7 7 7 7776 e i Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier not been described to date. Zoeae of Chionoecetes speciescarapace, and the setation of the abdominal somite 1. are similar in general morphology and setation, so that Rice (1980) felt that for the subfamily Oregoninae, distinguishing larvae can be difficult. A comparison of including the genera Hyas, Oregonia, and Chionoecetes , larval characteristics of Chionoecetes species (Tables 1-2) the posterior process on the scaphognathite in the first reveals that the larvae of C. tanneri share many characters zoeae is easily distinguished from the remaining marginal with congeners. setae. This is confirmed for C. bairdi (Haynes, 1973) and C. Haynes (1973) compared the first zoeae of C. bair di andtanneri (present study), but does not always appear to be C. opilio in terms of the length of the posterolateral spines the case in C. opilio and C. japonicus. In C. opilio the of abdominal somites and the length of the abdominal posterior process was uniquely distinguished in the larvae somites. Haynes (1973) described that the spines of C. described by Kurata (1963); however , this was not the case bairdi partly covered neighboring segments, whereas those in the larvae described by Motoh (1973) in which all the of C. opilio barely overlapped with the neighboring marginal setae were found to be identical in shape. This segment. In a comparative study of zoeae of C. opilio and was also the case for C. japonicus in which Motoh (1976) C. japonicus, Motoh (1982) found that differences were noted that the marginal setae were identical in shape and negligible in the relative length of the posterolateral spineslacked a posterior process. of abdominal somites and the length of the abdominalSetation of the telson using the presence and absence of a somites. Consequently, these characteristics vary not only minute lateral spine on the telson fork also appears to be from species to species b ut also within a species.highly a variable character within a species and therefore of Wencker et al. (1982) found considerable variability of limited use to distinguish among species. For C. opilio spine lengths in the zoeae of C. bairdi and C. opilio zoeae, Kurata (1963) and Kon (1980) described two lateral collected in the southeastern Bering Sea, and consequently spines (one large and one minute spines) and one mid- used additional characteristics to identify them. Those dorsal spine on the telson fork, whereas Motoh (1973) characteristics were: length from tip of the dorsal spine tonoted only one large lateral spine and one mid-dorsal spine. the tip of the rostral spine (described as RDSL in the Haynes (1973) described C. bairdi zoeae as having two present study), the shape and relative length of the carapace lateral spines and one mid-dorsal spine whereas C. lateral spines, and size of the dorsolateral processes (knobs)japonicus zoeae were described as having only one lateral of the abdominal somite 2-3. spine and one mid-dorsal spine (Motoh, 1976; Konishi and In Chionoecetes zoeae, the carapace lateral spine is longMatsumoto, 2002) similar to what was found in this present and straight or curved ventrally in the first zoea, and isstudy for C. tanneri zoeae. shorter and straight in the second zoea. Wencker et al. For the larval characteristics that are commonly used to (1982) noted that in C. bairdi zoeae, the lateral spines of thedescribe Chionoecetes species, the characters describing carapace are generally long and curved ventrally, whereas zoeae of C. tanneri overlap with many of those found in in C. opilio zoeae, the lateral spines are short and straight. similar larval stages of sympatric congeners. However, In C. opilio collected from Japanese waters, the dorsolateral when a comparison of the whole suite of characteristics is processes of zoeae I are longer (Kurata, 1963; Kuwatani etconducted for all described congeners, a pattern emerges by al., 1971; Kon, 1980) than those collected elsewhere. They which the larvae can be keyed to species. This information are also shorter in C. bairdi (Haynes, 1973), C. japonicus can be used to dev elo p dichoto mous keys for various larval (Motoh, 1976; Konishi and Matsumoto, 2002), and C.stages of Chionoecetes species. tanneri (the present study). These findings indicate that the size and shape of the carapace lateral spine in Chionoecetes ACKLOWEDGMENTS zoeae is too variable to be used as an identifying character. We extend special thanks to Mr. G. Workman and crews of CCGS The shape of the carapace lateral spines and the W . E. Ricker for collecting the ovigerous female crab. The work dorsolateral process for C. tanneri zoeae are similar to C. has been partially supported by The Basic Science Research bairdi zoeae, but they are different in size (RDSL). First Grant of Ministry of the Education, Korea. zoeae of C. tanneri (RDSL, 5.3-5.7 mm) are larger than those of C. bairdi, C. opilio, and C. japonicus. REFERENCES The previous studies often omitted detailed description of some important characteristics of Chionoecetes larvae. Aikawa H (1937) Further notes on brachyuran larvae. Rec. In addition to the larval characteristics noted above from Oceanogr. Works, Japan, 9: 87-162. previous work, the following features are important to Garth JS (1958) Brachyura of the Pacific coast of America. describe: the shape of the marginal setae on the scaphonathite Oxyrhyncha. Allan Hacock Pacific Expedition 21. 854 p. of the maxilla in the first zoea, the number of spines on the Hart JFL (1982) Crabs and their relatives of British Columbia. telson fork, the number of submarginal setae on the British Columbia Provincial Museum Handbook No. 4. 68 ANIMAL CELLS AND SYSTEMS Vol. 13 No. 1 Larv al Dev elopment of the Groov ed T anner Crab Victoria, BC 267 p. o p il io and C. japonicus reared in the laboratory . Univ . Alaska Sea Grant, Fairbanks, AK pp 119-136. Haynes E (1973) Description of prezoea and stage I zoea of Chinoecetes bair di and C. opilio (Oxyrhyncha, Oregoniinae).Park W and Shirley TC (2008) Variations of abundance and hatch Fish Bull 71: 769-775. timing of Dungeness crab larvae in southeastern Alaska: implications for climate effect. Animal Cells and Systems 12: Jamieson GS (1990) Development of a fishery for Chi ono ecete s 2 87- 29 5. ta nneri on the continental slope off British Columbia: management considerations. In: Proceedings of the Park W, Douglas DC, and Shirley TC (2007) North to Alaska: Symposium King and Tanner crabs, Melteff BR (ed.). Univ.evidence for conveyor belt transport of Dungeness crab larvae Alaska Sea Grant Fairbanks, AK. pp 587-592 along the west coast of the United S tates and Canada. Li mnol O ceano gr 52: 248-256. Jamieson GS, Heritage GD, and Noakes DN (1990) Life history characters of Chionoecetes tanneri off British Columbia. In: Pereyra WT (1966) Bathymetric and seasonal distribution, and Proceedings of the Symposium King and Tanner crabs, reproduction of adult tanner crabs, Chionoecetes tanneri Melteff BR (ed.). Univ . Alaska Sea Grant Fairbanks, AK. ppRathbun (Brachyura: Majidae), off the northern Oregon coast. 15 3- 162. D eep- S ea R e s 13: 1185-1205. Jewett SC and Haight RE (1977) Descriptions of megalopa of Pereyra WT (1968) Distribution of juvenile tanner crabs snow crab, Chionoecetes bairdi (Majidae, Subfamily (Chonoecetes tanneri) Rathbun, life history model, and Or ego n in ae) . Fish Bull 76: 459-463.fisheries management. Pr oc Na tl Sh ell fish A ssoc 58: 66-70. Kon T (1980) Studies on the life history of the Zuwai crab, Pereyra WT (1972) Bathymetric and seasonal abundance and Chionoecetes opilio (O. Fabricius). Spec Publ Sado Mar Biol general ecology of the tanner crab, Chionoecetes tanneri St, Nigata Univ Ser 2: 1-64. Rathbun (Brachyura: Majidae) of f the northern Oregon coast. In: Columbia River estuary and adjacent ocean waters: Konishi K and Matsumoto R (2002) The complete larval Bioenvironmental studies, Pruter A T and Alverson DL (eds.). development of Chionoecetes japonicus under laboratory Univ Washington Press, Seattle, WA pp 538-582. conditions. In: Crabs in Cold Water Regions: Biology, Management, and Economics, Paul AJ, Dawe EW , Elner R, Rice AL (1980) Crab zoeal morphology and its bearing on the Jamieson GS, Kruse GH, Otto RS, Sainte-Marie E, Shirley classification of the Brachyura. Trans Zool Soc London 35: T C , and W o o dby D (ed s.) . Univ . Al aska Sea Gr ant , F a ir bank s, 2 71- 42 4. AK. pp 135-146. Schmitt WL (1921) The marine decapod Crustacea of California Kurata H (1963) Larvae of Decapoda Crustacea of Hokkaido. 2. with special reference to the decapod Crustacea collected by Majidae (Pisinae). Bull Hokkaido Reg Fish Res Lab 27: 25- the United States Bureau of Fisheries steamer “Albatross” in 31 . connection with the biological survey of San Francisco Bay during the year 1912-1913. Univ Calif Publ Zool 23: 1-470. Kuwatani Y, Wakui T, and Nakanishi T (1971) Studies on the larvae and the post-larvae of a tanner crab, Chi ono ecete s W e ncker DL, In cze LS , and Armstron g DA (19 82) Distin guishing opilio elongatus Rathbun. I. On the protozoeal larvae. Bu llbetween Ch ion o ecet es ba ir di and C. opilio zoeae collected in Hokkaido Reg Fish Res Lab 37: 32-40. the southeast Bering Sea, Univ Alaska Sea Grant, Fairbanks, AK pp 119-136. Lough RG (1975) Dynamics of crab larvae (Anomura, Brachyura) off the central Oregon coast, 1969-1971. PhD W orkman GD, Phillips AC, Scurrah FE, and Boutillier JA (2002) Diss, Oregon State Univ., Corvallis, OR 299 pp.A new fishery for grooved Tanner Crab (Chionocetes tanneri ) off the coast of British Columbia, Canada, In: Paul AJ, Dawe Motoh H (1973) Laboratory-reared zoea and megalopae of Zuwai EG , Elner R, Jamieson GS, Kruse GH, Otto RS, Sainte-Marie crab from the Sea of Japan. Bull Japan Soc Sci Fish 39: 1223- B, Shirley TC, Woodby D (eds.). Crabs in Cold Water 12 30. Regions: Biology, Management, and Economics. University Motoh H (1976) The larval stages of benizuwai-gani, of Alaska Fairbanks, Alaska Sea Grant College Program. AK- Chionoecetes japonicus Rathbun reared in the laboratory. Bu ll SG-02-01, pp 439-456. Japan Soc Sci Fish 4 2 : 53 3- 542 . Motoh H (1982) The larval stages of the genus Ch ion o ecet es : C. [Received January 30, 2009; accepted February 16, 2009] ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 69
http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.pngAnimal Cells and SystemsTaylor & Francishttp://www.deepdyve.com/lp/taylor-francis/larval-development-of-the-grooved-tanner-crab-chionoecetes-tanneri-FeGlbhjz3j
Larval development of the grooved tanner crab, chionoecetes tanneri rathbun, 1893 (decapoda: Brachyura: Majidae) described from the laboratory‐reared specimens
Larval development of the grooved tanner crab, chionoecetes tanneri rathbun, 1893 (decapoda: Brachyura: Majidae) described from the laboratory‐reared specimens
Abstract
Abstract This paper documents the defining morphological characteristics of the larval stages of Chionoecetes tanneri Rathbun, 1893, the grooved Tanner crab, from specimens reared in the laboratory. Chionoecetes tanneri larval stages include two zoeae and one megalopa. The first zoea is characterized by: six setae on the posterior margin of the carapace; postero‐lateral spines on abdominal somites 3 and 4, extending beyond the posterior margin of adjacent somites and bearing 9–10...
Animal Cells and Systems 13: 59-69, 2009 Larval Development of the Grooved T anner Crab, Chionoecetes tanneri Rathbun, 1893 (Decapoda: Brachyura: Majidae) Described from the Laboratory- reared Specimens 1 2 2 Sung Y un Hong, W ongyu Park *, R. Ian Perry , and James A. Boutillier Department of Marine Biology , Pukyong National University , Pusan 608-737, Korea; Department of Marine Biotechnology , Soonchunhyang University , Asan 336-745, Kor ea; Department of Fisheries and Oceans, Pacific Biological Station, Nanaimo, B.C. V9T 6N7, Canada Abstra ct: This p aper document s the defining morphological INTRODUCTION characteristics of the larval stages of Chionoecetes tanneri Rathbun, 1893, the grooved Tanner crab, from specimens Chionoecetes tanneri Rathbun, 1893, the grooved Tanner reared in the laboratory . Chionoecetes tanneri larval stages crab, is a majid crab ranging in distribution from Kamtchatka, include two zoeae and one megalopa. The first zoea is Russia to Cortez Bank, Mexico (Schmitt, 1921; Garth, characterized by: six setae on the posterior margin of the 1 958; Ha r t , 1982 ) . Chionoecetes tanneri has been considere d carapace; postero-lateral spines on abdominal somites 3 and 4, extending beyond the posterior margin of adjacent for commercial exploitation in at least some parts of its somites and bearing 9-10 spinnules; 12 plumose setae and range, including the offshore waters of British Columbia, one stout distal plumose seta present on the margin of the Canada (Pereyra, 1966, 1968, 1972; Jamieson, 1990; scaphognathite of the maxilla; and one fused lateral spine Jamieson et al., 1990; Workman et al., 2002), thereby and one articulated dorso-medial spine on each fork of the providing part of the motivation for this study. Their telson. The second zoea is characterized by: 9 setae on the expected distribution is along the continental slope at postero-lateral margin of the carapace; a serrated mandible depths of 458-1784 m (Hart, 1982) and they do not appear molar; a mandibular palp bud; 25-26 plumose setae on the margin of the scaphognathite of the maxilla; pereiopods with to occur in coastal waters (Jamieson, 1990). They have well-developed gills and buds; and four pairs of stout setae been observed to occur in fine sediment habitats (Pereyra, on the posterior margin of the telson. For the megalopal 1966; Hart, 1982), and in aquaria have been observed to stage, the distinguishing characteristics include: a rostral feed on brittle stars and pieces of fish. In Oregon waters, spine equal in length to the supraorbital spine; six setae on they have been observed to release eggs, moult and mate the exopod of the uropod; and a single spine on the ischium during early spring, with reproduction taking place on an of the second pereiopod. This study allows C. t anneri larvae to be distinguished from the larvae of known sympatric annual cycle (Pereyra, 1966). However, considering their congeners. This information provides a basic taxonomic tool wide distribution range, they may have various reproductive for researchers in fisheries management and zooplankton schedules throughout the distribution range, as indicated in ecology who are addressing issues related to trophic Dungeness crab in the west coast of North America (Park et interactions, metapopulation dynamics and ecosystem al., 2007; Park and Shirley, 2008). Largely because of impacts in the evolving marine resource management identification prob le ms, larvae of g rooved T anner crab have strategies in the North Pacific, and those related to not been observed, but are assumed to occur near surface C h io noece te s species in particular. waters during late spring and summer; newly settled Key words: larval development, Chionoecetes tanneri , mega- benthic juveniles were observed below the depths of adult lopa, zoea, grooved tanner crab crabs (Pereyra, 1966). Chionoecetes contains five species: the Snow crab, C. * T o whom correspondence should be addressed. opilio; the Red queen crab, C. japonicus ; the Tanner crab, T el: +82-41-530-1282; Fax: +82-41-530-1638 E-mail address: pwg09@hotmail.com C. bairdi ; the grooved Tanner crab, C. tanneri ; and the ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 59 Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier triangle Tanner crab, C. angulatus. Of these, the larvae of formalin an d dissected in 10 0% ethylene glycol. The larvae C. opilio and C. japonicus have been described from were dissected using a M5 Wild binocular microscope. Japanese waters (Aikawa, 1937; Kurata, 1963; Kon, 1980; Appendages were illustrated using a M1 1 W ild microscope Konishi and Matsumoto, 2002), and those of C. opilio and equipped with a drawing tube. Measurements of key C. bairdi have been described from Alaskan waters morphological characteristics included: dorsal spine length (Haynes, 1973; Jewett and Haight, 1977). Larvae of C. (DSL); rostral spine length (RSL); distance between tips of tanneri were illustrated from plankton samples taken in dorsal and rostral spines (RDSL); lateral spine length Oregon waters (Lough, 1975); however, the larvae of C. (LSL); distance between tips of lateral spines (LLSL); tanneri have been difficult to distinguish from its sympatric carapace length as measured from the posterior margin of congeners because of the similarity and overlap of a the eye socket to the mid-dorsal posterior margin of the number of morphological characters. W encker et al. (1982) carapace for zoeae and from the tip of the rostrum to mid- reported finding some rare and unknown Chionoecetes dorsal posterior margin of the carapace for megalopa (CL). larvae of deep-dwelling Chionoecetes species from the A min i mu m of fi v e sp e ci me ns we re ex amin ed a n d me a sure d regions overlaying the co ntinental slope of the south eastern for each larval stage. Specimens used for the research have Bering Sea. Considering the deep-dwelling habit ranges of been deposited in the Invertebrate Laboratory, Pukyong C. tanneri and its larval characteristics, which did not National University, Pusan, Korea as PKNU-INVERT con form to th e published characteristics of C. bair di and C.number 02-01. opilio, it was postulated that the unknown Chionoecetes larvae may have been the larvae of C. tanneri or some other RE SU L T S deepwater Chionoecetes species. The objective of the present study is to describe the zoeae The larval development of C. tanneri Rathbun comprised and megalopa of C. tanneri reared in the laboratory and to two zoeal an d one megalopal stages. No meg alopa mo ulted compare them with previously described larvae of knownto the first crab stage. sympatric congeners. This study is intended to provide a taxonomic tool for researchers studying zooplankton inFirst zoea (Fig. 1) general and Chionoecetes species in particular to address questions of trophic interactions, metapopulation dynamics Size: CL=1.29 mm (1.20-1.38); DSL=2.40 mm (2.25- and ecosystem impacts in the evolving marine resource2.58); RSL=1.90mm (1.73-2.03); management strategies in the North Pacific.RDSL, 5.73mm (5.48-6.08); LSL=1.44mm (1.35-1.53). Duration: mean 30.7 (range 26-53) days. MA TERIALS AND METHODSColor: dark vermillion or dark crimson, chromatophores on abdominal somites 2-4, precardiac region and frontal st An ovigerous female C. tanneri was collected during a base of dorsal spine of the carapace, protopodite of 1 deep water assessment survey aboard the CCGS W. E.maxilliped, and mandibles. Ricker on 11 February 2000 from a crab pot fished at a Carapace (Fig. 1A, B). Dorsal, rostral, and lateral spines depth of about 400 m near Brooks Peninsula, Vancouver present; dorsal and rostral spines armed with numerous o o Island, British Columbia, Canada (128 00'W, 50 1 0 'N). In spinules on distal third; dorsal spine slightly longer than situ water temperature was around 5 C. The female was rostral spine; rostral spine slightly posteriorly curved; transported to the laboratory and maintained in a 400 L lateral spines, almost half as long as dorsal spine, armed aquarium at a constant temperature (8.8 C) until the larvae with numerous spinules, at right angle to carapace, curved hatched. slightly downward; a minute frontal knob between eyes; a Hundreds of larvae hatched from the female on 16 pair of antero-median and a pair of minute postero-dorsal February 2000. Of these, a single larva was transferred for setae; postero-lateral margin fringed with six submarginal rearing into each of one hundred 250 mL jars. An additional setae; margin with one plumose anterior seta and five 150 larvae were reared for mass culture in a 20 L jar. The posterior setae; lateral margin with two deep indentations; mass culture samples were used for drawing and descriptions. one just posterior to eye, one on the middle of the lateral Newly hatched Artemia nauplii were fed to the larvae.margin. Water temperature during the larval rearing ranged from Eyes (Fig. 1A, B). Tag eyes sessile onto the end of the o o o 8.8C through 10.4C (mean 9.6C); salinity from 26.0‰ tocarapace. 31 .0 ‰ (mean 28.9‰). W ater in each jar was changed daily Antennule (Fig. 1C). Uniramous; with 2 setae and 2 after examining the condition of the larvae, followed byaesthtascs. feeding with Artemia nauplii. Antenna (Fig. 1D). Biramous ; spinous process (protopod) Exuviae and specimens were preserved in buffered 5% elongate, subequal in length to rostral spine; armed with 60 ANIMAL CELLS AND SYSTEMS Vol. 13 No. 1 Larv al Dev elopment of the Groov ed T anner Crab numerous spinules; endopod rudimentary, a minutedistal segment. dendrical; exopod slender; less than one-third length of the Antenna (Fig. 2D). Endopod as an elongate bud, one- spinous process; two subterminal spines near acute tip. quarter the length of the spinous process (protopodite); Mandible (Fig. 1E). Incisor and molar processes well-exopod about one-half the length of spinous process. developed; palp absent. Mandible (Fig. 2E). Molar serrations, truncated and Maxillulae (Fig. 1F). Coxal endite with 2+6 setae; basial fused; both mandibles with a median indentation; endite with three fused plumodenticulate cuspidate setae, amandibular palp bud present. setal bud on the distal corner, and four setae; endopod 2- Maxillule (Fig. 2F). Basial endite with four fused segmented with 1, 2+4 plumodenticulate setae. plumodenticulate cuspidate setae and five setae; plumose Maxilla (Fig. 1G). Coxal endite, bilobed with 4 + 4 setae;exopod seta on outer margin; otherwise unchanged. basial endite, weakly bilobed with 5+5 setae; endopod, Maxilla (Fig. 2G). Basial endite with 5+6 setae; exopod bilobed with 3 + 3 setae; exopod (scaphognathite) margin(scaphognathite) margin with 25-26 plumose setae. with 12 plumose setae and one distal stout plumose seta. First maxilliped (Fig. 2H). Coxa without epipod bud; First maxilliped (Fig. 1H). Coxa with a denticulate seta; basis with an epipod bud and nine medial denticulate setae, basis with nine medial denticulate setae arranged 2, 2, 3, 2; arranged 2, 2, 3, 2; endopod setation unchanged; exopod endopod 5-segmented with 3, 2, 1, 2, 5 (1 subterminal + 4with six plumose natatory setae; otherwise unchanged. terminal) setae; exopod with four terminal plumose Second maxilliped (Fig. 2I). Basis with five medial natatory setae. denticulate setae; exopod with six plumose natatory setae; Second maxilliped (Fig. 1I). Coxa without setae; basisotherwise unchanged. with 4 medial denticulate setae arranged 1, 1, 1, 1; endopodThird maxilliped (Fig. 2J). Biramous epipod; with gill. 3-segmented with 1, 1, 5 (2 subterminal+3 terminal) setae;Pereiopods (Fig. 2K). Well-developed; with gill buds. exopod with 4 terminal plumose natatory setae. Abdomen (Fig. 2A, L). Six somites plus telson; somites Third maxilliped (Fig. 1I). Present; biramous. 2-6 with a pair of mid-dorsal setae and a pair of pleopod Pereiopods (Fig. 1I). Present; chela bilobed. buds; the posterolateral spine on somites 3-5 naked and Abdomen (Fig. 1A, J). Five somites plus telson; somite 1 barely extended beyond the posterior margin of the with (or rarely without) a pair of mid-dorsal setae; somites adjacent somites; somite 6 with a pair of short 2-3 with a pair of small, subequal, dorsolateral processes;posterolateral spine. somites 2-5 with a pair of posterodorsal setae. Somites 3-5 Telson (Fig. 2A, B, L). Telson fork with an articulated with a pair of posterolateral spines; the spines on somites 4-lateral spine and an articulate dorsomedial spine; furca with 5, armed with 9 or 10 spinules and extended beyond four pairs of stout posterior marginal setae of which the posterior margins of adjacent somites; spine on somite 5,innermost pair is much smaller than the others. n a k ed . T elson (Fig. 1A, J). T elson fork long and flexed distallyMegalopa (Figs. 3 and 4) with a fused lateral spine and an articulate dorsomedial spine; mid-posterior margin deeply concave with a medialSize: CL=3.25mm (2.90-3.55). arch; furca with three pairs of stout posterior marginal Duration: eleven megalopae obtained from 23 second setae. zoeae; survived for 45 days; no megalopas molted to first crab. Second zoea (Fig. 2) Carapace (Fig. 3A, B). Longer than wide; rostrum directed anteroventrally with a pair of stout anterolateral Size: CL=1.83 mm (1.25-2.05); DSL=2.59 mm (2.25- processes; protogastric region with a pair of stout dorsal 3.05 .); RSL = 1.96 mm (1.83-2.13); RDSL = 5 .5 1 mm (5.35- spines; branchial and metagastric regions elevated; 5.70); LSL=0.96mm (0.83-1.05).intestinal region with a pair of stout dorsal processes; lateral Duration: 30.6 (29-34) days branchial regions with a pair of lateral protuberances; six Color: Similar to first zoea.pairs of setae on protogastric and intestinal regions. Carapace (Fig. 2A, B). Lateral spine almost a quarter ofEyes (Fig. 3A). Stalked. dorsal spine; ventral margin with an indentation posterior to Antennule (Fig. 3C). Peduncle 3-segmented with 0, 2, base of antenna; posterolateral margin fringed with nine 3 + 1 setae; endopod 2-segmented with 0,4 setae; flagellum submarginal or marginal setae.3-segmented 0, 4+1, 8 aesthetascs, and setae. Eyes (Fig. 2A, B). Stalked. Antenna (Fig. 3D). Peduncle 4-segmented with 0,0,1,4 Antennule (Fig. 2C). 3-segmented; proximal segmentsetae; flagellum 5-segmented with 0,3,4,0,3 setae. with a minute seta on the basal outer margin, one aesthetasc Mandible (Fig. 3E). Cutting edges well-developed, not on the penultimate segment, and eight aesthetascs on the clearly divided into incisor and molar processes; palp 2- ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 61 Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier Fi g. 1. T he gr oov ed T anner c rab, Chonoec etes t anner i Rat hbun, 1893, fir s t z oea. A, l ateral v i ew ( ab); B, f ront al v i ew ( ab); C, ant ennul e ( ae) ; D, antenna (ac); E, mandibles (ae); F , maxillule (ae); G , maxilla (ae); H, first maxilliped (ac); I, second maxilliped (ac); J, ab domen and telson (ad). Scale bars with dif ferent sizes at the lef t bottom indicate 0.5 mm. segmented with 0, 12 setae. endite with 4 + 20 setae; endopod 2-segmented with 0, 3 Maxillule (Fig. 3F). Coxal endite with 14 setae; basialsetae. 62 ANIMAL CELLS AND SYSTEMS Vol. 13 No. 1 Larv al Dev elopment of the Groov ed T anner Crab Fi g. 2. The grooved Tanner crab, C hon oec etes t anner i Rat h bun, 18 93, s e c ond z oea. A, l a teral vi ew ( a b) ; B, fr on t a l v i ew (ab); C, a n t enn ul e (ae); D, antenna (ad); E, mandibles (ae); F , maxillule (ae); G , maxilla (ae); H, first maxilliped (ac); I, second maxi lliped (ac); J, thi rd maxi lliped (ae); K, pereiopod buds (ad); L, abdomen and telson (ac). Scale bars with dif ferent sizes at the left bottom indicate 0.5 mm. Maxilla (Fig. 3G). Coxal endite bilobed with 1 1+6 setae; process with 14 long setae; coxal and basial endites with basial endite bilobed with 9+10 setae; endopod with a numerous setae; endopod not distinctly segmented, distal naked process and 6 marginal plumose setae; scaphognathite part flattened, with two setae; exopod (scaphognathite) 2- fringed with 64-70 plumose setae.segmented with 1, 4 plumose setae. First maxilliped (Fig. 4A). Epipod, a long blade-like Second maxilliped (Fig. 4B). Coxa with one epipod and ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 63 Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier F ig. 3. T h e gr ooved T anner c r a b , Chonoecet es t a n ner i Rat h bun, 1893, megalop a. A, lat e r a l v i ew ( ab); B, dors a l v i ew (ab), C, antennule ( ad); D , a nt en na (ad); E, m andible (ae); F , maxill u l e (af ) ; G , maxilla (ae); H, later a l v i ew of abdomen (ac ) . Sc ale b a r s wi th dif f er ent s iz es at the lef t bottom ndicate 0.5 mm. one gill bud, epipod with five long setae; endopod 5- exopod 2-segmented, distal segment with six terminal segmen te d with 0,1,1,4,9 setae; exopod 2-segmen ted , distalplumose setae. segment with 0,4 plumose setae. Pereiopods (Fig. 4D-H). Pereiopods 1-3 with gills; all Third maxilliped (Fig. 4C). Coxa with 11 setae and one segmented and sparsely spinous; pereiopod 2 with a spine epipod with 30 long setae; basis/ischium with 3/25-30on basipodite. spines; merus with 10-11 spines; carpus with six spines;Abdomen (Figs. 3A, B, H). Six somites with 2, 5, 4, 4, 1, propodus with 7-8 spines; dactylus with seven spines; 1 pairs of surface setae; posterolateral mar gin of so mite 2-5 64 ANIMAL CELLS AND SYSTEMS Vol. 13 No. 1 Larv al Dev elopment of the Groov ed T anner Crab Fig. 4. The grooved T anner crab, Chonoecetes tanneri Rathbun, 1893, megalopa. A, first maxilliped (ac); B, second maxilliped (ac); C, third maxilliped (ac); D-H, pereiopod 1-5 (ab); I, pleopod on abdominal somite 1 (ac). Scale bars with different sizes at the left bottom indicate 0.5mm . broadly truncated.Telson (Fig. 3A, B, 4J): Longer than wide; naked margin. Pleopods (Figs. 3A, H, 4I). Present on abdominal somite 2-5; endopod, a simple rod-like bud with 2 or 3 curved DISCUSSION processes; exopods of pleopod 2-5 with 6,7,6,7 plumose natatory setae. Of the five Chionoecetes species in the Pacific Ocean, the Uropods (=pleopod on somite 6) (Figs. 3A, B, 4J): larvae of C. opilio, C. japonicus , and C. bairdi have been Uniramous with six plumose natatory setae. described, while those of C. tanneri and C. angulatus have ANIMAL CELLS AND SYSTEMS Vo l. 13 No. 1 65 6 S 6 u T able 1. Comparison of the larval characteristics of Chionoecetes first zoeae C. opilo elongatus C. opilio C. opilio C. opilio C. japonicus C. japonicus C. bairdi C. tanneri W (Konishi and (Jewett and Haight, (Kurata, 1963) (Haynes, 1973) (Motoh, 1973) (Kon, 1980) (Motoh, 1976) (The present study) Matsumoto, 2002)1977) Carap ace Carapace length (CL) 1.8-2.0 mm 0.54 (0.45-0.73) mm 1.83 (1.25-2.02) mm Dorsal spine length (DSL) 2.3 mm 5.33 (5.04-5.78) mm 2.59 (2.25-3.05) mm Rostral spine length (RSL) 1.8-2.0 mm 1.96 (1.83-2.13) mm Dorsal-rostral spine length (DRSL) 4.5-4.9 mm 4.6-5.4 mm 4.6-5.2 mm 5.21 (4.85-5.52) mm 1.24 (1.11-1.39) mm 4.17 (3.96-4.55) mm 5.51 ( 5.35-5.70) mm Lateral spine length (LSL) 1.3 mm 0.96 (0.83-1.05) mm Lateral-Lateral spine length (LLSL) 3.3-3.6 mm 3.3-3.5 mm 3.69 (3.46-3.93) mm 2.73 (2.52-2.79) mm Posterolateral margin seta 3* 3** 5 6 6 Antennule3+22+152+13+22+2 Antenna i Exopod/Spinous process <1/2 1/4 Endopodite A minute denticle minute tubercle A small protunerance A minute denticle Maxillule Coxal endite setae 7 7 7778 Basial endite setae 6 or 7 7 7 7 7+1 7+1 Endopod setae 1,6 1,6 1,6 1,6 1,6 1,6 Maxilla Coxal endite setae 4+4 4+4 4+4 4+4 4+4 4+4 Basial endite setae 5+(4 or 5) 5+5 5 (or 6)+5 (or 4) 5+6 5+5 5+5 Endopod setae 3+3 3+3 3+3 3+3 3+3 3+3 Scaphognathite setae 15+1 (10 or 11)*(2 or 3) (11-13)+1 12 or 13 12 or 13 11 (or rarely 12)+1 12+1 First maxilliped Basial setation 2+2+3 2+2+3+3 2+2+3+3 2+2+3+3 3+3+2+2 2+2+3+2 N Endopod setation 3,2,1,2,5 3,2,1,2,5 3,2,1,2,5 3,2,1,2,5 3,2,1,2,5 2,1,1,2,5 Second maxilliped Basial setation 1+1+1+1 1+1+1+1 1+1+1+1 1+1+1+1 1+1+1+1 1+1+1+1 Endopod setation 1,1,5 1,1,5 1,1,5 1,1,5 1,1,5 1,1,5 1 A Table 1. Continued C. opilo elongatus C. opilio C. opilio C. opilio C. japonicus C. japonicus C. bairdi C. tanneri (Konishi and (Jewett and Haight, (Kurata, 1963) (Haynes, 1973) (Motoh, 1973) (Kon, 1980) (Motoh, 1976) (The present study) Matsumoto, 2002)1977) Abdomen YSomite 1-5 or somite Dorsal paired setae on somites Somite 1-5 Somite 2-5 Somite 1-5 2-5 Dorsolateral process on somite 2 Straight Straight Slightly curved Slightly curved Straight l Dorsolateral spine on somite 3 Straight Slightly curved Slightly curved Straight Barely extending Extending a little Extending beyond beyond the bit beyond the Extending beyond Extending beyond Extending beyond Extending beyond the posterior Posterolateral spine on somite 3posterior posterior the posterior the posterior margine the posterior the posterior margine margine of somite margine of margine of margine of somite 4 of somite 4 margine of somite 4 of somite 4 somite 4 somite 4 Not quite Extending beyond E x ten d in g a litt l e Extending beyond Extending beyond Extending beyond Extending beyond reaching the the posterior bi t b e yon d th e Posterolateral spine on somite 4 the posterior the posterior margine the posterior the posterior margine poterior margine margine of somite posterior margine margine of somite 5 of somite 5 margine of somite 5 of somite 5 of somite 5 5 of so mite 5 Numerous Covered by 1 or 1 spinules 7 Spinules Spinules Spinules Few spinules 9-10 spinules spinules Posterolateral spine on somite 5SmoothSmooth Telson Lateral fused spine 1 1 1 1 1 1 1 Minute secondary spine 1 1 1 1 None Dorsomedial articulated spine 1 1 1 1 1 1 1 *Motoh (1973, Fig. 1B), **Motoh (1976, Fig. 1C) Table 2. Comparison of the larval characteristics of Chionoecetes megalopae v C. opilo elongatus C. opilio C. opilio C. japonicus C. japonicus C. bairdi C. tanneri (Konishi and Matsumoto, (Jewett and Haight, (Kurata, 1963) (Motoh, 1973) (Kon, 1980) (Motoh, 1976) (The present study) p 2002)1977) m Carapace Carapace length (CL) 3.6 mm 2.9-3.3 mm 3.0 mm 2.89-3.37 mm 2.83-3.26 mm 3.12-3.48 mm 2.90-3.55 mm Equal to theLonger than the Equal to the Equal to the Equal to the Three times longer Equal to the e Rostral spine lengthsupra-orbitalsupra-orbital supra-orbital supra-orbitalsupra-orbital than the supra-orbital supra-orbital c spinespinespinespinespinespinespine A minute lateral spine on AbsentAbsentAbsentAbsentAbsentPresentAbsent pterygostomial-branchial region Spine on basipodite of preiopds 0,1,1,1,01,1,1,1,0?0,1,1,1,00,1,1,1,01,1,1,1,00,1,0,0,0 1,2,3,4,5 Uropod Exopod seate 7 7 7 7776 e i Sung Y un Hong, Wongy u P ark, R. Ian Perry , and James A. B outillier not been described to date. Zoeae of Chionoecetes speciescarapace, and the setation of the abdominal somite 1. are similar in general morphology and setation, so that Rice (1980) felt that for the subfamily Oregoninae, distinguishing larvae can be difficult. A comparison of including the genera Hyas, Oregonia, and Chionoecetes , larval characteristics of Chionoecetes species (Tables 1-2) the posterior process on the scaphognathite in the first reveals that the larvae of C. tanneri share many characters zoeae is easily distinguished from the remaining marginal with congeners. setae. This is confirmed for C. bairdi (Haynes, 1973) and C. Haynes (1973) compared the first zoeae of C. bair di andtanneri (present study), but does not always appear to be C. opilio in terms of the length of the posterolateral spines the case in C. opilio and C. japonicus. In C. opilio the of abdominal somites and the length of the abdominal posterior process was uniquely distinguished in the larvae somites. Haynes (1973) described that the spines of C. described by Kurata (1963); however , this was not the case bairdi partly covered neighboring segments, whereas those in the larvae described by Motoh (1973) in which all the of C. opilio barely overlapped with the neighboring marginal setae were found to be identical in shape. This segment. In a comparative study of zoeae of C. opilio and was also the case for C. japonicus in which Motoh (1976) C. japonicus, Motoh (1982) found that differences were noted that the marginal setae were identical in shape and negligible in the relative length of the posterolateral spineslacked a posterior process. of abdominal somites and the length of the abdominalSetation of the telson using the presence and absence of a somites. Consequently, these characteristics vary not only minute lateral spine on the telson fork also appears to be from species to species b ut also within a species.highly a variable character within a species and therefore of Wencker et al. (1982) found considerable variability of limited use to distinguish among species. For C. opilio spine lengths in the zoeae of C. bairdi and C. opilio zoeae, Kurata (1963) and Kon (1980) described two lateral collected in the southeastern Bering Sea, and consequently spines (one large and one minute spines) and one mid- used additional characteristics to identify them. Those dorsal spine on the telson fork, whereas Motoh (1973) characteristics were: length from tip of the dorsal spine tonoted only one large lateral spine and one mid-dorsal spine. the tip of the rostral spine (described as RDSL in the Haynes (1973) described C. bairdi zoeae as having two present study), the shape and relative length of the carapace lateral spines and one mid-dorsal spine whereas C. lateral spines, and size of the dorsolateral processes (knobs)japonicus zoeae were described as having only one lateral of the abdominal somite 2-3. spine and one mid-dorsal spine (Motoh, 1976; Konishi and In Chionoecetes zoeae, the carapace lateral spine is longMatsumoto, 2002) similar to what was found in this present and straight or curved ventrally in the first zoea, and isstudy for C. tanneri zoeae. shorter and straight in the second zoea. Wencker et al. For the larval characteristics that are commonly used to (1982) noted that in C. bairdi zoeae, the lateral spines of thedescribe Chionoecetes species, the characters describing carapace are generally long and curved ventrally, whereas zoeae of C. tanneri overlap with many of those found in in C. opilio zoeae, the lateral spines are short and straight. similar larval stages of sympatric congeners. However, In C. opilio collected from Japanese waters, the dorsolateral when a comparison of the whole suite of characteristics is processes of zoeae I are longer (Kurata, 1963; Kuwatani etconducted for all described congeners, a pattern emerges by al., 1971; Kon, 1980) than those collected elsewhere. They which the larvae can be keyed to species. This information are also shorter in C. bairdi (Haynes, 1973), C. japonicus can be used to dev elo p dichoto mous keys for various larval (Motoh, 1976; Konishi and Matsumoto, 2002), and C.stages of Chionoecetes species. tanneri (the present study). These findings indicate that the size and shape of the carapace lateral spine in Chionoecetes ACKLOWEDGMENTS zoeae is too variable to be used as an identifying character. We extend special thanks to Mr. G. Workman and crews of CCGS The shape of the carapace lateral spines and the W . E. Ricker for collecting the ovigerous female crab. The work dorsolateral process for C. tanneri zoeae are similar to C. has been partially supported by The Basic Science Research bairdi zoeae, but they are different in size (RDSL). First Grant of Ministry of the Education, Korea. zoeae of C. tanneri (RDSL, 5.3-5.7 mm) are larger than those of C. bairdi, C. opilio, and C. japonicus. REFERENCES The previous studies often omitted detailed description of some important characteristics of Chionoecetes larvae. Aikawa H (1937) Further notes on brachyuran larvae. Rec. In addition to the larval characteristics noted above from Oceanogr. 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