16 Apri11996

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):53-69. 1996 Systematics and distribution of the Calocarides (Crustacea: : )

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.

Abstract.- The nine currently recognized of Calocarides are re- viewed, and two new species are described: C. capensis from the south-east Atlantic Ocean off the Cape of Good Hope, and C. macphersoni from the Atlantic Ocean off Namibia. Species of the genus are particularly characterized by the possession of dentate median, submedian, and lateral carinae of the carapace, weakly pigmented or unpigmented eyes, and robust, quite heavily sclerotized, tuberculate and/or spinose first pereopods.

The approximately 100 described species seum of Natural History, Smithsonian ofAxiidae show their greatest diversity in Institution. shallow tropical and subtropical seas, with several genera confined to the tropics, e.g., Systematic Account Coralaxius Kensley & Gore, 1982, Parax- Family Axiidae iopsis De Man, 1905, Paraxius Bate, 1888, Genus Calocarides Wollebaek, 1908 Scytoleptus Gerstaecker, 1856. A few gen- Calocaris (Calocarides) Wollebaek, 1908: era, however, have colonized colder and 3,23. deeper waters and show considerable diver- Axiopsis (Calocarides).-De Man, 1925:1, sity and broad geographic range, e.g., Cal- 2,6,67, 71.-Balss, 1957:1579. ocarides Wollebaek, 1908, and Calocarides.-Bouvier, 1940:97.-Sakai & Bate, 1888, the latter an inquiline of hex- de Saint Laurent, 1989:4, 78.-Poore, actinellid sponges. Calocarides is geo- 1994:98. graphically widespread in continental shelf- Euconaxius [laps. cal. for Eiconaxius Bate, slope regions, and contains cold-water in- 1888] Trybom, 1904:384. habiting, soft-bottom burrowing species, Type species.-By subsequent designa- some of which may be associated with tion by Sakai & de Saint Laurent, 1989, Eu- zones of upwelling. The genus is reviewed conaxius coronatus Trybom, 1904. Type lo- here, two new species are described, and a cality: SkagelTak, North Sea, 230-500 m. number of taxonomic and distributional Gender: Masculine. problems noted, relati ng to species previ- Diagnosis.-Gonochoristic. Carapace ously included in the genus. having small supraocular spine present, as Abbreviations: cl-carapace length; part of lateral rostral series; post-cervical HUMZ-Laboratory of Marine Zoology, carina low, rounded, barely discernible in Hokkaido University; ICMB-Instituto de some species, lacking spines; rostrum at Ciencias del Mar, Barcelona; MNHN-Mu- same level as anterior carapace, margins seum National d'Histoire Naturelle, Paris; armed; median carina dentate; submedian SAM-South African Museum, Cape carina dentate; lateral carina dentate. Town; UBZM-Zoological Museum, Uni- Eye reaching to about midlength of ros- versity of Bergen; USNM-National Mu- trum, cornea weakly pigmented or unpig- Table I.-Branchial formula for Calocarides coron- Calocarides soyoi (Yokoya, 1933). Off atus. r = reduced. northern Japan, 138-270 m. Calocarides spinulicauda (Rathbun, 1902). Off Bodega Head, California, 113 m; Pu- get Sound, 70-256 m; British Columbia, Exopod 91-128 m. Epipod Podobranch r - Calocarides vigila Sakai, 1992. New Zea- Arthrobranch 222 2 land, 74-146 m. Pleurobranch Setobranch Taxonomic remarks.-The species Cal- ocaris lev Zarenkov, 1989, described from a hydrothermal vent at 2000-2036 m in the Gulf of California, is almost certainly a spe- mented; eyestalk longer than eye. Antennal cies of Calocarides. The proportions and scaphocerite elongate. Maxilla 2, scaphog- spination of the first chelipeds bear more nathite having spinulose flagellum. Maxil- resemblance to C. rostriserratus from lipeds 1-3 with exopods and epipods. Pe- Chile, Peru, and Panama, than to C. quin- reopod I, chelipeds asymmetrical, some- queseriatus from southern California. Giv- what sexually dimorphic. Propodi and dac- en that this species was taken from greater tyli of pereopods 3-5 slender, not depth than ei ther of the two abovemen- expanded, dactyli simple. Pleopodal rami tioned species, its relative isolation of the slender elongate; appendix interna present Gulf of California, and the lack of males, it on pleopods 2-5. Pleopod I 0 absent; pleo- seems prudent to await further material be- pod 1

V;g?

Maxilliped 3, merus with 4 spines on eral surface with few scattered small tuber- posterior margin; carpus with single small cles and ridge near posterior margin, ante- distal spine. Pereopod 1, larger cheliped, is- rior margin with about 11 spines; fixed fin- chium with single strong distal spine; merus ger and dactylus as in larger cheliped. with strong spine at midlength of posterior Pereopod 2, ischium with few small spines margin, few small tubercles in proximal and strong distal spine on posterior margin; half, anterior margin with 5 distal spines; merus with 3 widely spaced spines on pos- carpus with few low tubercles on anterior terior margin. Pleopod 1 consisting of sin- margin and few small tubercles on lateral gle slender elongate ramus. Pleopods 2-5 surface; propodal palm slightly longer than each with appendix interna on mesial mar- fingers, outer surface with tiny scattered gin of endopod. Lateral uropodal ramus acute tubercles and strong ridge near pos- with single small spine on outer margin; su- terior margin reaching almost to tip of fixed ture bearing 4-8 irregular spines; mesial ra- finger, anterior margin with about 11 strong mus with 4 spines on middorsal ridge. spines, cutting margin of finger finely Remarks.-See 'Remarks' after Calocar- toothed, with large proximal tubercle; dac- ides macphersoni. tylus with few tubercles on anterior margin, Etymology.- The specific name derives cutting edge finely toothed, with proximal from the type locality, the Cape of Good tubercle. Smaller cheliped, ischium with Hope. strong distal spine; merus with strong spine at midlength of posterior margin, anterior Calocarides coronatus (Trybom, 1904) margin with 3 distal spines; carpus with few Euconaxius coronatus Trybom, 1904:384- tubercles on anterior margin and lateral sur- 390, pI. 20, figs. 1-10,13,14, pI. 21, figs. face; propodal palm longer than fingers, lat- 1-8. . , " - . •~. " .

'~'

Fig. 2. Calocarides capensis, new species: A, Carapace in lateral view, scale = 5 mm; B, Anterior carapace in dorsal view; C, Abdomen in lateral view; D, Telson in dorsal view; E, Pleopod I, female; F, Pleopod 2, female; G, Pereopod I, smaller cheliped; H, Pereopod I, larger cheliped; I, Maxilliped 3; J, Pereopod 2. Calocaris (Calocarides) coronatus.-Wol- Antennal scaphocerite reaching distal lebaek, 1908:3, 5, 13, 23. margin of peduncle article 4. Maxilliped 3, Axiopsis (Calocarides) coronatus.-De merus with 4 spines on posterior margin, 3 Man, 1925:6, 67, 71. more distal spines large; carpus with small Calocarides coronatus.-Bjorck, 1913:7.- distal spine on posterior margin. Pereopod Balss, 1925:209.-Poulsen, 1940:208, 1, larger cheliped: ischium with row of 216, fig. 4.-Bouvier, 1940:97.-Chris- acute tubercles on posterior margin, distal- tiansen, 1955: 1--4; 1972:40, fig. 45.- most a large spine; merus with posterior Elofsson, 1959:3-9, figs. 1-20.-Bratte- surface expanding distally, mesial and lat- gard, 1966:45-52.-Sakai & de Saint eral margin defined by row of small acute Laurent, 1989:4,80, 101, fig. 20. tubercles, mesial margin with strong spine Euconaxius crassipes Trybom, 1904:390- at about midlength, anterior margin bearing 391, pI. 20, figs. 11, 12. 1 strong spine and few blunt tubercles; car- Calocaris (Calocarides) crassipes.- Wol- pus bearing small tubercles dorsodistally; lebaek, 1908:3,5-13, 17,23, pI. 1-7. fingers slightly more than half length of Axiopsis (Calocarides) crassipes.-De propodal palm, latter with lateral and most Man, 1925:6, 67, 71. of mesial surface tuberculate, fixed finger with triangular tooth at about midlength of Material examined.-UBZM 50400, (; cutting edge; fingers widely gaping, dacty- 0 cl 14.8 mm, Sorfjord, Norway, 60 11'N, lus downcurved, strongly tuberculate, with 6°34'E, 589-600 m.-UBZM 49598, 4 (; 1 or 2 stronger tubercles on cutting edge. cl 13.5, 13.9, 15.5, 15.5 mm, Kvinnherad- Smaller cheliped: ischium, merus, and car- fjord, Norway, 59°56'N, 5°45'E, 512 m.- pus as in larger cheliped; propodal palm UBZM 49595, <;> cl 14.0 mm, Kvinnherad- with lateral and mesial surface tuberculate, fjord, Norway, 59°59'N, 5°53'E, 660-670 tubercles along anterior margin forming m.-UBZM 50461, ovigerous <;> cl 14.4 spines, fingers about 213 length of palm, mm, Osafjord, Norway, 60040'N, 6°55'E, fixed finger with blunt proximal triangular 300.-UBZM 49589, Alfjord, Norway, tooth followed by regularly spaced small 59°41'N, 5°34'E, 450-465 m.-UBZM, teeth; dactylus with row of spines along an- ovigerous <;> cl 16.1 mm, Fusafjord, Nor- terior margin, cutting edge with regularly 0 way, 60 12'N, 5°45'E, 240 m.-UBZM spaced small teeth. Pereopod 2, ischium 49591, (; cl 15.1, <;> cl 15.2 mm, Bomla- and basis each with single posterodistal fjord, Norway, 59°40'N, 5°22'E, 360 m. spine; merus with 3 evenly spaced spines Distribution.-From Malangenfjord, on posterior margin. Lateral uropodal ra- northern Norway, to southwestern Sweden, mus with about 18 spines along suture, 2 80-700 m. small distal spines on lateral margin; mesial Diagnosis.-Carapace surface smooth. ramus with single distolateral spine, dorsal Rostrum with 4 pairs of lateral teeth, con- ridge bearing 5 or 6 spines. tinuous with lateral carina bearing 3 teeth; median carina with 1 tooth anterior to tu- Calocarides macphersoni, new species bercle; submedian carina bearing 2 teeth. Fig. 3 Abdominal pleuron 1 ventrally narrowly Calastacus longispinis.-Macpherson, rounded; pleuron 2 broadly rounded; pleura 1983:45, fig. 26; 1991:405, non Mc- 3-5 rounded, with 1 or 2 small teeth on Ardle, 190 I. anterior margin. Telson with 5 or 6 lateral teeth, 2 mobile spines at posterolateral an- Material examined.-Holotype, USNM gle; posterior margin evenly convex, with 243561, (; cl 20.4 mm, RlV Benguela II sta median tooth; I or 2 small spines proxi- 66, off Namibia, 20055'S, 12°23'E, 475- modorsally on rounded ridge. 480 m, 9 Sep 1980.-Paratypes, USNM Fig. 3. Calocarides macphersoni, new species: A, Lateral view, scale = 5 mm; B, Telson and right uropod in dorsal view; C, Pereopod 2; D, Maxilliped 3; E, Pereopod I, larger cheliped; F, Pereopod 1, smaller cheliped.

243562, 0" cl 17.8 mm, RJV Benguela II sta us sp.-Paratypes, ICMB, RJV Benguela II 66, off Namibia, 20055'S, 12°23'E, 475- sta P-66, 2 0" cl 20.3 mm, one damaged, 480 m, 9 Sep 1980.-Paratype, ICMB, RJ off Namibia, 20055'S, 12°23'E, 475--480 V Benguela II sta P-37, 0" cl 21.5 mm, off m.-Paratypes, ICMB, RJV Benguela IV Namibia, 21°33'S, 12°39'E, 380-390 m, sta P-95, 2 0" cl 21.0 mm, 21.2 mm, off from stomach of scorpaenid fish Helicolen- Namibia, 20042'S, 12°14'E, 410--417 m, one specimen from stomach of ray Raja small tubercles along posterolateral margin, confundens, one from stomach of scorpaen- anterior margin with row of ilTegularly id fish Helicolenus sp. large and small spines and tubercles; outer Diagnosis.-Rostrum with 2-3 pairs of and upper surface of carpus fairly densely lateral teeth; median carina with single tooth tuberculate; propodal palm about Ys longer anterior to tubercle, entire posterior to tu- than fingers, posterolateral carina feebly tu- bercle; submedian carina poorly defined, berculate, extending almost to apex of fixed with 1 or 2 teeth; lateral carina with 3-5 finger, outer and upper surfaces tuberculate, teeth posterior to supraocular. Carapace sur- anterior margin with double row of large face smooth. First abdominal pleuron ven- and small spines and tubercles; cutting mar- trally narrowly rounded, pleura 2-6 broadly gins of fixed propodal finger and dactylus rounded, small spine on anterior margin of only slightly excavate proximally, remain- pleura 3-6. Telson with 5 small fixed spines der of margin bearing row of small evenly- in posterior half of lateral margin, 3-4 small spaced teeth. Pereopod 2, posterior margin mobile spines mesial to posteriormost lateral with row of small spines and single strong spine; posterior margin gently convex be- distal tooth; posterior margin of merus with tween lateral spine and median spine; dorsal 2-4 spines. Lateral uropodal ramus with 2 surface with one pair of small fixed spines. or 3 small distal spines on lateral margin, Maxilliped 3, posterior margin of ischi- single mobile spine mesial to distal most, um unarmed; merus with 3-4 strong distal transverse suture armed with row of 13-15 spines and 2-3 tiny more proximal spines small spines; mesial ramus with single dis- on posterior margin; carpus with small pos- tal spine, dorsal ridge with irregular row of terodistal spine. Pereopod 1: larger chela, about 8 small spines. ischium with row of small denticles and Remarks.-From McArdle's (1901) de- single strong spine along posteromesial scription, and Alcock & McArdle's illustra- margin, small distal spine on anterior mar- tions (1902, pI. 57, figs. 2, 2a), and De gin; merus with posteromesial surface tu- Man's (1925:118) key, the Bay of Bengal berculate, having single strong tooth at species Calastacus longispinis lacks spines about midlength, anterior margin with on the anterior median carina, has five about 5 small spines and several small tu- spines on the free lateral margins of the ros- bercles; outer and upper surface of carpus trum, and three spines on the submedian ca- evenly tuberculate; propodal palm about Ys rina. The antennal scaphocerite reaches al- longer than fingers, entire surface evenly most to the end of peduncle article 4. The and fairly densely tuberculate, posterior single female lacked both chelipeds of pe- margin with tuberculate carina extending reo pod 1. Given that the Namibian material onto fixed finger, anterior margin with row has a strong spine anterior to the tubercle of somewhat larger tubercles; fingers prox- on the median carina, and that the antennal imally widely gaping, distally meeting, scaphocerite reaches only to the distal third fixed finger with proximal half of cutting of peduncle article 4, it seems unlikely that edge having few low rounded tubercles, the Atlantic and Indian Ocean specimens distal half straight, bearing numerous small are the same species, especially considering rounded teeth; surface of dactylus tubercu- the considerable geographical separation. late, cutting edge as in fixed propodal fin- The species described above as Calocar- ger. Smaller chela, ischium with few small ides capensis, differs from C. macphersoni spines and single strong spine on posterior in having more slender and less granular margin, single distal spine on anterior mar- chelipeds of pereopod 1, with relatively gin; merus with row of small tubercles more elongate spines on the anterior margin along posteromesial margin with 1 or 2 of the propodal palm; in having fewer but strong spines at about midlength, row of stronger distal spines on the anterior margin of the merus of pereopod 1; in having far carcinologist Dr. Enrique Macpherson of fewer spines on the suture of the latera] ra- Barcelona, Spain, who made the material mus of the uropod; and in having a more from Namibia availab]e for study. elongate te]son. Major differences between C. coronatus· Calocarides quinqueseriatus and C. macphersoni are seen in the size (el (Rathbun, 1902) ovigerous <;? 13.5-16.1 mm vs. c] ovigerous Figs. 4, 5 <;? 17.8-25.0 mm resp.), and in the more Calastacus quinqueseriatus Rathbun, 1902: spinose upper margin of the merus in C. 887.-Schmitt, 1921:113, fig. 76.- macphersoni. Balss, 1925:209. The species described as Axius (Neaxius) Calocaris (Calastacus) quinqueseriata.- laevis by Bouvier (19] 5) from 698 m off De Man, 1925:8, 1] 8. Cape Bojador, Spanish Sahara, West Africa, Calocarides quinqueseriatus.-Sakai & de (holotype, MNHN, <;? cI 5.1 mm) was syn- Saint Laurent, 1989:79, 103. onymized with C. coronatus by Sakai & de Saint Laurent (1989:81, fig. 21). The latter Material examined.-Syntypes, USNM authors illustrated the single damaged spec- 25240, 9 0 el 15.5-22.7, 2 ovigerous <;? el imen of C. laevis, the caption of their illus- 15.4, 18.2,3 <;? cI 17.5-19.4, 19.4, RlV Al- tration erroneously referring to Eiconaxius batross sta 3196, San Luis Obispo Bay, laevis. Given the damaged condition of the California, green mud, 366 m.-USNM specimen, and the fact that it lacks spines 28321,50, el13.5-21.6, 1 ovigerous <;? el on the submedian carina, has an antennal 16.9 mm, 9 <;? c] 15.1-20.5, RlV Albatross scaphocerite shorter than in C. coronatus, sta 2909, Santa Barbara Channel, Califor- has pereopod 1 lacking the fine tubercula- nia, 375 m, green mud.-USNM 152525, tion of the propodus and the anterior spi- 22 0 c] 18.9-26.0, 9 ovigerous <;? 20.0- nation of the merus of C. coronatus, and a 23.3, II <;? el 21.0-23.0, RlV Albatross sta telson longer and narrower than in C. co- 4436, San Migue] Island, California, 483- ronatus, it seems unlikely that the West Af- 496 m, green mud.-USNM 28325, 16 0, rican species is synonymous with C. coron- el 11.3-] 8.0, 1 ovigerous <;? cl 24.0, 7 <;? el atus, or indeed, that it is a species of Cal- 10.2-19.2, RlV Albatross sta 3195, San ocarides. From the figure of Calocarides Luis Obispo Bay, California, 461 m, green rudolfi (Zarenkov, 1989, fig. 3) from An- mud.-USNM 28236,5 0 c] 14.8-24.0,4 gola, severa] differences in spination sug- <;? c] ]3.0-22.5, RIV Albatross sta 3198, off gests that it differs from C. macphersoni. In Point Conception, California, 508 m, green C. rudolji, the merus of maxilliped 3 bears mud.-USNM 28327,7 0 el 12.9-20.5, 3 more elongate and numerous spines, and ovigerous <;? cI ]9.0-20.0,6 <;? cl 10.3-19.5, the merus of both chelae of pereopod 1 has RlV Albatross sta 3199, Santa Barbara larger and more numerous anterodistal and Channel, California, 426 m, green mud.- posterodistal spines than in the Namibian USNM 28232, 0 cl 18.0, 2 <;? cl 21.0, 21.9 species; the carpus of pereopod 2 bears a RlV Albatross sta 3187, off Point Sur, Cal- posterodistal spine, and the telson bears two ifornia, 545 m, brown and grey ooze.- pairs of fixed dorsa] spines, while C. mac- USNM 28320, 0 cl 20.3, <;? c] 17.2, RIV phersoni lacks a carpal spine on pereopod Albatross sta 2892, off Point Conception, 2 and the telson bears a single pair of fixed California, 520 m, yellow mud.-USNM spines. Posterolateral mobile spines on the 28528, 2 0, cI 18.1, 24.], RIV Albatross te]son were not illustrated for the Ango]an sta 3200, Santa Barbara Channel, Califor- species; C. macphersoni bears four such nia, 485 m, green mud.-USNM 28319, 1 spines on each side. o cI 19.0, I <;? cl 12.1, RIV Albatross sta Etymology.- The species is named for 2891, off Point Conception, California, 426 Fig. 4. Calocarides quinqueseriatus: A, Lateral view, scale = 10 mm; B, Pereopod 1, larger cheliped; C, Pereopod I, smaller cheliped; D, Telson and right uropod in dorsal view; E, Pleopod 2, male; F, Pleopod 3, male; G, Pleopod I, female; H, Pleopod 2, female. m, brown ooze.-USNM 120119, « cl m, green mud.-USNM 53021, « cl 16.2, 18.2, RN Commando sta, mouth of Colum- RN Albatross sta 4425, off San Nicolas Is- bia River, Oregon, 732 m.-USNM 28324, land, California, 2013 m, green mud, fine o cl 17.5, « cl 15.0, RN Albatross sta sand, globigerina ooze.-USNM 155735, 3193, off San Simeon Bay, California, 293 o cl 20.5, RlV Albatross sta 4387, off San m, green mud.-USNM 28329, ovigerous Diego, California, 1830 m, green mud.- « cl 19.5, RJV Albatross sta 3201, Santa USNM 243350, ovigerous « damaged, RJ Barbara Channel, California, 512 m.- V Commando sta, SW mouth of Columbia USNM 28322, 0 cl 17.4, RN Albatross sta River, Oregon, 1556 m.-USNM 243351, 2979, off Anacapa Island, California, 710 o cl 18.4, RN Commando sta, SW mouth -r ~ --. -r -r --~ ~-r---7-..--7-r~ --rr ~ ..., --1 o E

~~ ...~ .. ~.... )

G

Fig. 5. Calocarides quinqueseriatus, variation in rostral and anterior carapace spination: A, Syntype, ovi- gerous female, San Luis Obispo Bay, California, cl 15A mm; B, Syntype, ovigerous female, San Luis Obispo Bay, California, cl 18.2 mm; C, Ovigerous female, San Miguel Is., California, cl 22.0 mm; D, Ovigerous female, San Miguel Is., California, cl 23.3; E, Female, Baja California, cl 12.0 mm; F, Female, Baja California, 20.7 mm; G, Male, Columbia River mouth, Oregon, 18.2 mm; H, Syntype, male, San Luis Obispo Bay, California, cl 19.1 mm; I, Syntype, male, San Luis Obispo Bay, California, cl 20.9 mm; J, Male, San Miguel Is., California, 24.2 mm; K, Male, San Miguel Is., California, cl 26.0 mm; L, Male, Columbia River, Oregon, cl 27.2 mm.

of Columbia River, Oregon, 1373 m.- California, 445 m, green mud.-USNM USNM 243352, damaged, R/V Comman- 243355, 2 0' cl 16.5, 2 cl 13.8-15.0, R/ do sta, SW mouth of Columbia River, Or- V Albatross sta 4351, NE of Point Lorna, egon, 1556 m.-USNM 243353, 0' cl 17.3, California, 774 m, soft green mud. 2 damaged, RJV Commando sta, SW Diagnosis.-Carapace smooth; 5-6 pairs mouth of Columbia River, Oregon, 1922 rostral teeth including small supraocular; m.-USNM 243354, damaged, R/V median carina with 1--4 spines anterior to Commando sta, SW mouth of Columbia tubercle, 1-2 posterior; 1-5 spines on sub- River, Oregon, 1556 m.-USNM 155737, median carina; 3-7 spines on lateral carina. cl 20.0, R/V Albatross sta 5675, San Abdominal pleuron 1 with ventral spine Cristobal Bay, Baja California, 520 m.- usually present; pleuron 2 ventrally round- USNM 155736,2 0' cl 19.8-21.0, RJV Al- ed; pleura 3-6 rounded with spine on an- batross sta 4433, off Santa Rosa Island, terior margin. Telson with lateral margin having 5-6 small spines; pair spines on Size ranges.-Male, maximum carapace dorsal surface. length 26.0 mm; ovigerous females, cara- Maxilliped 3, ischium with about 10 pace length 15.4-24.0 mm. small spines on posterior margin; merus Remarks.- There is considerable varia- with 5-6 small proximal spines, 3 large dis- tion in the spination of the anterior carapace tal spines on posterior margin. Pereopod 1, (Fig. 5) and the chelipeds of pereopod 1 of larger (broader) cheliped: ischium with 1 this species, often made more marked either strong distal and several small proximal by injuries to the chelae that have healed, spines on posterior margin; merus with in- or by loss and regeneration of the chelipeds, ner and outer row of tiny tubercles flanking resulting in a range of cheliped proportions posterior row of about 17 strong spines, and armature. outer surface distally with few scattered The majority of specimens have been tiny tubercles, upper (anterior) margin with taken from soft bottom sediments, especial- irregular row of spines, distal spines largest; ly from green mud. carpus with outer surface and distal margin The species recorded as Calastacus quin- bearing scattered acute tubercles; upper queseriatus from 1150 m off Japan (Kob- propodal palm subequal in length to fingers, jakova 1937: 142, pI. 2, fig. 8) appears to posterior margin of palm and fixed finger a have the first pereopod chelae relatively denticulate carina, upper (anterior) margin shorter, more robust, and less granular than with about 14 strong spines, mesial and lat- in true C. quinqueseriatus. It thus seems un- eral surfaces of palm fairly densely granu- likely that the Japanese species is conspe- lar/tuberculate, fixed finger with broad tu- cific with the California/Oregon species. bercle on proximal cutting edge, distally denticulate; dactylus cutting edge with Calocarides rostriserratus proximal tubercle, distally denticulate, up- Andrade & Baez, 1977 per margin with several proximal spines. Fig. 6 Pereopod I, smaller (narrower) cheliped: is- Calastacus rostriserratus Andrade & Baez, chium, merus, and carpus as in larger che- 1977:65, fig. 1. liped; upper propodal palm subequal in Calocarides quinqueseriatus.-Sakai & de length to fingers, tuberculation as in larger Saint Laurent, 1989:78, 79, 103, non cheliped, cutting edge of fixed finger with Rathbun, 1902. few small tubercles proximally, remainder of edge finely denticulate; dactylus, cutting Material examined.-USNM 173366, 4 edge finely denticulate, with about 2 spines 0' cl 15.3 mm, 15.3 mm, 16.3 mm, 17.0 on upper margin. Pereopod 2, ischium with mm, 2 S! cl 14.5 mm, 15.9 mm, off Tum- 6 small spines on posterior margin; merus bes, Peru, 3°33.8'S, 81°01.4'E, 576 m, 28 with 5 small proximal spines, single strong Aug I979.-USNM 243392, S! cl 16.1 mm, distal spine. Pleopod 1 in female a slender RlV Gilliss sta GS-22, Gulf of Panama, elongate ramus; pleopod 2 in male with 7°28'N, 79°12'W, 825 m, 18 Jan 1972. rod-shaped appendix masculina bearing dis- Diagnosis.-Carapace smooth; rostrum tal setae, slightly shorter than appendix in- with 5-6 pairs of lateral teeth including terna; pleopods 2-5 in female, 3-5 in male small supraocular; 2 median carina teeth with appendix interna at about proximal plus tubercle; 4-5 teeth on submedian ca- third of mesial margin of endopod. Uropod rina; 4-6 teeth on lateral carina. Abdominal with lateral ramus having 2 small distal pleura 2-5 ventrally rounded. Telson with spines on lateral margin, single mobile sub- 2-3 pairs small lateral spines, 2 small distal marginal spine; about 14 spines along su- mobile spines at posterolateral angle, 2 ture; mesial ramus with single distolateral pairs spines on dorsal surface. spine, 4 spines on dorsal surface. Maxilliped 3, ischium with 5 small prox- ~~ ~s

Fig. 6. Calocarides roslriserralUs: A, Lateral view, scale = 5 mm; B, Anterior carapace in dorsal view; C, Telson and right uropod in dorsal view; D, Pereopod I, larger cheliped; E, Pereopod I, smaller cheliped; F, Pleopod I, female; G, Pleopod 2, female; H, Pleopod 2, male; T, Pleopod 3, male. imal teeth on posterior margin, distal mar- palm 1.2 times length of fingers, bearing gin entire; merus with 2 strong distal teeth row of 8 strong spines, lower border con- on posterior margin. Pereopod 1, larger sisting of flattened distally narrowed ridge (broader) cheliped, ischium with about 5 having dentate margins, medial and lateral small proximal and single strong distal surfaces bearing moderately dense scattered spine on posterior margin; merus with 5 conical tubercles, cutting edge of fixed fin- distal spines on upper (anterior) margin, ger having rounded proximal cusp; dactylus posterior margin with single row of spines with upper margin bearing about 8 strong of varying length, those at about midlength spines, cutting edge having rounded proxi- longest; carpus with several (6-7) small mal cusps, distally finely denticulate. Pereo- spines on upper surface; upper propodal pod 1, smaller (narrower) cheliped, ischium with 2-3 small proximal and single strong Axiopsis (Axiopsis) soyoi.-Sakai, 1987: distal spine on posterior margin; merus with 303. 5-6 strong distal spines on anterior (upper) Calocarides soyoi.-Sakai & de Saint Lau- margin, single row of spines along posterior rent, 1989:4, 83, 103.-Kensley & Ko- (lower) margin, those at about midlength mai, 1992:81, fig. 1. longest; carpus with 8-10 spines on upper surface; upper propodal palm about 1.2 Material examined.-USNM 243563, 2 times length of fingers, bearing irregular (j> cl 12.0 mm, 15.0 mm, ovigerous (j> cI row of abou t 12 spines, lower border 17.9 mm, off Fukushima Prefecture, Japan, formed by flattened ridge with dentate mar- 3r04'N, 141°31.7'E, 270 m.-USNM gins, inner and outer surfaces bearing scat- 243564, <3cI 14.2 mm, ovigerous (j> cI 18.8 tered acute conical tubercles; dactylus with mm, (j> cl 11.8 mm, off Hachinohe, Aomori several small spines on upper surface, cut- Prefecture, Japan.-HUMZ-C990, 3 (j> cl ting edge more or less evenly denticulate. 11.3 mm, 14.1 mm, 13.0 mm, off Hachi- Pereopod 2, basis with single spine on pos- nohe, Aomori Prefecture, Japan, 42°52.6'N, terior surface; ischium with proximal and 145°22.2'E, 152 m.-HUMZ-C994, (j> cI one or 2 stronger distal spine on posterior 16.9 mm, off Fukushima Prefecture, Japan, margin; merus with 3 or 4 spines in mid- 37°17.3'N, 141°21.4'E, 141 m.-HUMZ- region of posterior margin. Pleopod 1 in fe- C998, 3 <3cI 12.4 mm, 15.0 mm, 15.3 mm, male a slender elongate ramus; pleopod 2 off Fukushima Prefecture, Japan, 37°04'N, in male with rod-shaped appendix mascu- 141°31.7'E, 270 m. lina bearing distal setae, noticeably shorter Remarks.- This species was recently re- than appendix interna; pleopods 2-5 in fe- described and figured by Kensley & Komai male, 3-5 in male with appendix intema at (1992). about proximal third of mesial margin of endopod. Uropod with lateral ramus having Calocarides spinulicauda 2-3 small lateral spines plus distal submar- (Rathbun, 1902) ginal mobile spine; about 13 spines along Fig. 7 suture; mesial ramus with single distal mar- ginal spine, 4-5 spines on upper surface. Axius spinulicauda Rathbun, 1902:886; Remarks.-In spite of Sakai & de Saint 1904:149, fig. 90. Laurent's (1989:80) statement that C. ros- Axiopsis spinulicauda.-Schmitt, 1921: 111, triserratus is probably a synonym of C. fig. 74.-Butler, 1961 :60, figs. 1, 2, pI. 1. quinqueseriatus, Calocarides rostriserratus Axiopsis (Axiopsis) spinulicauda.-De differs from C. quinqueseriatus, its geo- Man, 1925:6, 67, 69. graphically nearest congener, in several eas- spinulicaudus.-Sakai & de ily seen features: in having two pairs of Saint Laurent, 1989:4, 66, 103. dorsal telsonic spines (one pair in C. quin- Material examined.-Holotype, USNM queseriatus), and two or three lateral tel- 25239, (j> cI 15.5 mm, R/V Albatross sta sonic spines (five or six in C. quinqueser- 3172, off Bodega Head, California, 113 m. iatus); and in having the chelae of pereopod I broader and less elongate, and with spines Diagnosis.-Carapace surface smooth; only in the distal half of the anterior margin rostrum with 5 (left) and 6 (right) lateral of the merus (full anterior margin of merus teeth, including supraocular; median carina bearing spines in C. quinqueseriatus). barely reaching onto base of rostrum, with 3 spines anterior of tubercle, entire poste- Calocarides soyoi (Yokoya, 1933) rior to tubercle; postcervical median carina Axius soyoi Yokoya, 1933:49, fig. 25.- obsolete except close to posterior margin of Horikoshi et aI., 1982:30, 33, 38, 39, 120, carapace; submedian carina short, entire; 145, 168. lateral carina entire posterior to supraocular Fig. 7. Calocarides spinulicauda: A, Carapace in lateral view, scale = 5 mm; B, Abdomen in lateral view; C, Anterior carapace in dorsal view; D, Telson and right uropod in dorsal view. spine of rostral series. First abdominal pleu- distal spines on posterior margin. Pereopod ron ventrally narrowed apically rounded 1, both chelipeds missing. Pereopod 2, me- with tiny denticle; pleura 2-6 broadly rus with 2 or 3 well separated spines on rounded. Telson with lateral margin bearing posterior margin. Lateral uropodal ramus larger fixed anterior spine, 4-5 small more with 7 small distal spines on lateral margin, posterior fixed spines; posterior margin transverse suture with row of 11-12 small evenly convex; submedian longitudinal spines; mesial ramus with 2 strong spines ridge bearing 3 fixed spines, pair of fixed on lateral margin, dorsal ridge with 7 or 8 anterior spines mesial to longitudinal ridge small fixed spines. present. Remarks.-Rathbun (1902), in her de- Maxilliped 3, ischium with 1 or 2 tiny scription of A. spinulicauda, noted that the denticles on posterior margin; merus with 3 chelae of the first pereopod were missing. Butler (1961) figured both chela of pereo- iidae).-Anales del Musec de Historia Natural pod 1, from material from British Colum- de Valparaiso 10:65-67. Balss, H. 1913. Diagnosen neuer ostasiatischer Ma- bia. While the lateral and mesial faces of cruren.-Zoologischen Anzeiger 62(5):234- the propodal palm do not appear to be tu- 239. berculate, as is characteristic of Calocari- ---. 1925. Macrura del' Deutschen Tiefsee-Expe- des, the spination of the dactyli, propodi, dition. I. Palinura, Astacura und Thalassinid- carpi, and meri are characteristic of this ge- ea.-Wissenschaftliche Ergebnisse del' Deutsch- en Tiefsee-Expedition auf dem Dampfer "Val- nus. Calocarides spinulicauda, however, divia" 1898-1899, 20(4): 189-216. disagrees with Calocarides in lacking spi- ---. 1957. Decapoda.-Dr. H. G. Bronn's Klassen nation on the lateral and submedian carinae. und Ordnungen des Tierreichs 5(1)7( 12): 1505- Sakai & de Saint Laurent (1989) placed this 1672. species in their new genus Acanthaxius, Barnard, K. H. 1950. Descriptive catalogue of South Af- which is characterized by a tendency to rican decapod Crustacea (Crabs and Shrimps).- Annals of the South African Museum 38: 1-837. having a double cornea, and a lack of pleu- Bate, C. S. 1888. Report on the Crustacea Macrura robranchs, neither of which are seen in C. collected by H.M.S. Challenger during the years spinulicauda. With only the female holo- 1873-76.-Report on the Scientific Results of type available for examination, the nature the Voyage of H.M.S. Challenger during the of the male pleopods and the first pereopods years 1873-76, Zoology 24:i-xc, 1-942. Bjorck, W. 1913. Decapoden aus dem Kattegatt und being uncertain, unambiguous generic aus dem Skagerak.-Arkiv fOr Zoologi 8(3): 1- placement is almost impossible. 12. Axius spinulicauda amurensis Kobjako- Bouvier, E. L. 1915. Thalassinides nouveaux captures va, 1937, (see also Derjugin & Kobjakova au large des c6tes Soudanaises par Ie "Talis- 1935:142), described from the Sea of Japan, man."-Bulletin du Museum National d'His- toire Naturelle, Paris 21(6):182-185. while clearly not a subspecies of the Cali- 1940. Decapodes Marcheurs.-Faune de fornian Calocarides spinulicauda Rathbun, France 37: 1-399. 1902, is difficult to place generically, given Brattegard, T. 1966. Ecological and biological notes the single small figure and the abbreviated on Calocarides coronatus (Crustacea Thalassi- description. nidea).-Sarsia 24:45-52. Butler, T. H. 1961. Records of decapod Crustacea from British Columbia.-Canadian Journal of Acknowledgments Zoology 39:59-62. Christiansen, E. N. 1955. Notes on Calocarides co- I thank Dr. Enrique Macpherson of the ronatus (Trybom) (Crustacea Decapoda).-As- Instituto Ciencias del Mar, Barcelona, Mrs. tarte [1112:1-5. Michelle van der Merwe of the South Af- Derjugin, K. M., & S. I. Kobjakova. 1935. Zur De- rican Museum, and Dr. A. Fosshagen of the kapodenfauna des Japanischen Meeres.-Zool- Zoological Museum, Bergen, for the loan ogischer Anzeiger 112(5/6):141-147. of material. The paper was considerably im- Elofsson, R. 1959. A new decapod larva referred to Calocarides coronatus (Trybom).-Universite- proved by the comments and suggestions of tet i Bergen, Arbok 1959, Naturvitenskapelig Dr. Rafael Lemaitre and anonymous re- rekke 7:1-10. viewers, to whom I am very grateful. Gerstaecker, A. 1856. Carcinologische Beitrage.-Ar- chiv fUr Naturgeschichte 22(1):101-162. Gurney, R. 1942. Larvae of Decapod Crustacea. Lon- don, Ray Society. 306 pp. Alcock, A., & A. F. McArdle. 1902. Illustrations of Horikoshi, M., T. Fujita, M. Takeda, M. Okiyama, S. the zoology of the Royal Indian Marine Survey Ohta, E. Tsuchida, & M. Yamamoto. 1982. ship Investigator, under the command of Com- Preliminary compilation of the results obtained mander T. H. Heming, RN. Crustacea, part 10, by the "Survey of Continental Shelf Bordering plates 56-67. Calcutta: Office of the Superin- Japan" carried out on board the S/S S6y6-Maru tendent of Government Printing, India. during 1923-1930. In M. Horikoshi, ed., Report Andrade, H., & P. Baez. 1977. Calasracus rostriser- for a part of the result of Grant-in-Aid for Co- raws n. sp. (Crustacea, Decapoda, Macrura, Ax- operative Research (A), Ministry of Education, Science and Culture: "Studies on Off-shore and Poulsen, E. M. 1940. On the occurrence of the Thal- Deep-sea Faunas in the West Pacific and Indian assinidea in Danish waters.-Videnskabelige Oceans" (Proj. no. 00534033). Ocean Research Medelelser, Copenhagen 104:207-239. Institute, University of Tokyo, 252 pp. Rathbun, M. J. 1902. Descriptions of new decapod Kensley, B. 1981. On the zoogeography of southern from the west coast of North Amer- African decapod Crustacea, with a distributional ica.-Proceedings of the United States National checklist of the species.-Smithsonian Contri- Museum, 24( 1272):885-905. butions to Zoology 338: 1-64. ---. 1904. Decapod crustaceans of the northwest ---. 1994. The genus Coralaxius redefined, with coast of North America.-Harriman Alaska Ex- the description of two new species (Crustacea: pedition 10:1-210. Decapoda: Axiidae).-Journal of Natural His- Sakai, K. 1987. Two new Thalassinidea (Crustacea: tory 28:813-828. Decapoda) from Japan, with the biogeograph- ---, & R. H. Gore. 1982. Coralaxius abelei, new ical distribution of the Japanese Thalassinid- genus and new species (Crustacea: Decapoda: ea.-Bulletin of Marine Science 41 (2):296-308. Thalassinidea: Axiidae): a coral inhabiting 1992. Axiid collections of the Zoological shrimp from the Florida Keys and the western Museum, Copenhagen, with the description of Caribbean Sea.-Proceedings of the Biological one new genus and six new species (Axiidae, Society of Washington 103(3):558-572. Thalassinidea, Crustacea).-Zoologica Scripta ---, & T Komai. 1992. Redescription of Calocar- 21(2):157-180. ides soyoi (Yokoya, 1933) from Japan (Crusta- ---, & M. de Saint Laurent. 1989. A check list of cea: Decapoda: Axiidae).-Proceedings of the Axiidae (Decapod a, Crustacea, Thalassinidea, Biological Society of Washington 105( I):81-85. Anomura), with remarks and in addition descrip- Kobjakova, S. \. 1937. Systematische Uebersicht der tions of one new subfamily, eleven new genera Dekapoden aus dem Ochotskischen und Japan- and two new species.-Naturalists 3:1-104. Schmitt, W. L. 1921. The Marine Decapod Crustacea ischen Meere.-15:93-154. [In Russian, with of California, with Special Reference to the German summary.] Decapod Crustacea Collected by the United Macpherson, E. 1983. Crustaceos Decapodos captu- States Bureau of Fisheries Steamer "Albatross" rados en las costas de Namibia.-Resultados in connection with the Biological Survey of San Expediciones Cientfficas (Sup!. Inv. Pesq.) II: Francisco Bay during the Years 1912-1913.- 3-79. University of California Publications in Zoolo- ---. 1991. Biogeography and community struc- gy, 23:1-470, pis. I-SO, 165 text figures. ture of the decapod fauna off Na- Smith, S. I. 1881. Preliminary notice of the Crustacea mibia (Southeast Atlantic).-Journal of Crus- dredged, in 64 to 325 fathoms, off the south tacean Biology 11(3):401-415. coast of New England, by the United States McArdle, A. F 190 I. Natural history notes from the Fish Commission in I880.-Proceedings of the Royal Indian Marine Survey Ship 'Investiga- United States National Museum 3:413-452. tor', Commander T H. Heming, R. N., com- Stebbing, T R. R. 1910. General catalogue of South manding. Series 3, No.5. An account of the African Crustacea (Part V of S. A. Crustacea, trawling operations during the surveying-season for the Marine Investigations in South Afri- of 1900-190 I.-Annals and Magazine of Nat- ca).-Annals of the South African Museum ural History (7)8:517-526. 6(4):281-593. Man, J. G. de. 1905. Diagnoses of new species of ma- Trybom, F 1904. Two new species of the genus Eu- crurous decapod Crustacea from the "Siboga-Ex- conaxius.-Arkiv for Zoologi 1(4):383-393. pedition".-Tijdschrift der Nederlandsche Dier- Wollebaek, A. 1908. Remarks on decapod crustaceans kundige Vereeniging (2)9(3/4):587-614. of the North Atlantic and the Norwegian Fiords ---. 1925. The Decapoda of the Siboga-Expedi- (I and II).-Bergens Museums Aarbog 12: 1-74. tion. Part VI. The Axiidae collected by the Si- Yokoya, y. 1933. On the distribution of decapod crus- bog a-Ex pedi tion .-S iboga- Ex ped itie monogra- taceans inhabiting the continental shelf around phie 39a5:1-127. Japan, chiefly based upon the materials collect- Poore, G. C. B. 1994. A phylogeny of the families of ed by S. S. S6y6-Maru, during the years 1923- Thalassinidea (Crustacea: Decapoda) with keys 1930.-Journal of the College of Agriculture, to families and genera.-Memoirs of the Mu- Tokyo Imperial University 12(1): 1-226. seum of Victoria 54:79-120. Zarenkov, N. A. 1989. Three new species of Calo- ---, & D. J. G. Griffin. 1979. The Thalassinidea caris Bell (Decapoda Axiidae).-Zoologiskii (Crustacea: Decapoda) of Australia.-Records Jhurnal 68(11):24-30. [In Russian, with English of the Australian Museum 32(6):217-321. summary.]