Biol ogy of Anomura II (A. Asakura,ed. ),C rustacean Research,Specia lNumber 6: 133-149,2006

Descriptions of color patterns in selected species of pagurid hermit crabs with sexual tubes (Decapoda: Anomura: Paguridae) from

Akira Asakura

Abstract.-In decapod crustaceans,color- guished by coloration of the ocular peduncles ation is often useful in taxonomic studies for and shield . separating species that are difficult to distin- In this paper,color patterns of eight species guish by morphological characters but are of pagurid hermit crabs in which males have reproductively isolated. In this paper,color pat・ asexual tube(s) are described on the basis of terns are described for eight pagurid species color photographs of live animals or specimens with sexual tubes; Catapaguroides cristimanus in which color pattern was well preserved. de Saint Laurent 1968 Turleania senticosa , , These information has become available dur- (McLaughlin and Haig 1996),Catapagurus hi- , ing various my studies on hermit crab fauna of rayamai (Asakura,2001) ,Catapagurus kosugei Japan. Since all of these species are small and (Asakura,2001) ,Nematopagurus gardineri morphologically similar to other congeners the Alcock,1905a ,Anapagurus japonicus Ortmann, , color information is useful for future taxonomic 1892,Boninpagurus acanthocheles Asakura & Tachikawa,2004 ,and Trichopagurus trichophthal- study of these species or genera,in particular, mus(Forest ,1954). These patterns are based on for comparative purpose. This is the first time color photographs of Ii ve animals or specimens in for these species to describe color pattern in de- which color patterns were well preserved. tails . In addition to examination of color pat- terns of the Japanese specimens,the speci- INTRODUCTION mens housed in the following museums were In decapod crustaceans,many examples are examined for morphological comparison; known for groups of species that are difficult Museum National d'Histoi re Naturelle,Paris to distinguish by morphological characters (MNHN); National Museum ofNatural History, but are reproductively isolated (Knowlton, Smithsonian ln stitution ,Washington D.C. 1986,1993). However,decapods are primar- (USNM). ily described and identified by morphological characters,which can be studied in preserved SYSTEMATlCS specimens,so that these problematic taxa might sometimes be overlooked.Many non- Catapaguroides cristimanus de Saint Laurent, morphological characters have been used in 1968 distinguishing such species,including karyol- Figs. 1,2A ,B ogy,life history,development ,mating behavior, Catapaguroides cri stimanus de Saint Laurent,1968: biochemical characters,molecular data and so 949,fi g. 30.一一 Asakura,2004 ・ 456,fi g. 1. on. Among them,color pattern is particularly important (Knowlton,1986) .For example , Materialexa mined .-IO ,SL = 1. 0mm , diogenid hermit crabs Calcinus gaimardii (H. 200 m ,dredge ,off Onga-zaki,Ishigaki lsland , Milne Edwards,1848) and Calcinus morgani Yaeyama Group,Ryukyu Islands,R 八1Yaeyama Rahayu & Forest,1999 ,are primarily distin- (cap. Koei Higa),21 Nov.2002 ,col l. Takeharu 134 A. Asakura

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Fig. 1. Catapaguroides cr istimanusd eSaint Laurent,196 8: o ,SL = 1.0 mm,Ishigaki-jima Island ,Y aeyama Group,R yukyu Islands ,CBM-ZC 8779. A ,left second pereopod,later al;B ,same ,dactyl ,propodus and carpus,mesial; C ,left third pereopod,latera l. Setae omitted in all figures. Scales = 1mm .

Kosuge,C8 恥4・ZC8779; 1 ♀,SL = l. 3mm , proximal angle of dorsal face of palm,latero ・ sledge net,off Kakeroma Island ,Amami distal angle of dorsal face of ca中 us,and mesial Islands,s t. ll ,R /V Toyoshio・maru,11 Dec. and lateral faces ventrally of,and dorsodistal 1995,col l. Susumu Ohtsuka,C8M-ZC 8780. angle of,merus. Type material examined (for morphologi- Left cheliped (Fig. 2A) generally white, cal comparison): holotype,o ,SL = 1.5 mm, with subdistal transverse faint orange band Siboga st. 260,90 m ,Key Islands,lndonesia ,5 present on dactyl and fixed finger,small brown 036.05'S,132 055'02E,16 Dec.1899 ,恥骨 >!HN patches present on mesioproximal angle of Pg;paratypes :2 <;>, SL = 1.3 ,1.3 mm,data dorsal face of palm,laterodistal angle of dorsal same as Pg. face of carpus,and mesial and lateral faces ven- trally of,and dorsodistal angle of,merus . Description 01color pattern (two days after Second and third pereopods (Figs. 1,28) fixation in 10% formalin solution).- Shield generally semitransparent white,with small (Fig.2A) white,with small brown areas on an- brown patches. Dactyls with several ,s mall terior margins between rostrum and lateral pro- patches dorsally.Propodi with some small jections ;ocular peduncles pale brown,darker patches dorsally ;lateral faces with distal and distally; ocular acicles whitish proximal small patches.Carpi each with small Antennular and antennal peduncles (Fig. proximal patch,and on third ,additional subdis- 2A) whitish. tal patch on lateral face; mesial faces eachwith Right cheliped (Fig. 2A) generally white, distal small patch.Meri ofthird each with distal with small brown patches present on mesio- small patch. Color pattems ofhermit crabs 135

Fig. 2. Catapaguroides cristimanus de Saint Laurent,1968 ・o ,SL = 1.0 mm,Ishigaki-jima Island,Yaeyama Group,Ryukyu Islands,CBM-ZC 8779. A ,anterior half of body and appendages,dorsal ;B ,second (upper) and third (l ower) pereopods,latera l. Turleania senticosa (McLaughlin & Haig,1996): ♂, SL = 1.6 mm, IshigalくiIsland,Yaeyama Group,Ryukyu Islands ,CBM-ZC 8782. C ,who lebody ,dorsa l;D ,right cheliped, mesial ;E ,left cheliped,mesial 136 A .Asakura

Remαrks.-See Asakura (2004) for mor- (Figs. 2C,3A) mottled yellow-orange and phological description of these same speci- white,with small darker orange areas near gas- mens. 1have no chance to examine coloration tric pits; ocular peduncles whitish,with distal of live animals. Colored patch巴s and areas in 0.4 -0.5and proxima1 0.2-0.3 orange; ocular these specimens are mostly orange or brown, acicles whitish,with tips orange. and these may be artificia l. In my experience, Antennu1ar peduncles (Figs. 2C,3A) gen- green or blue color in hermit crabs,especially erally whitish,each with distal ,medial ,and in small species,are changed into orange or proximal na汀 oworange bands on ultimate seg- brown immediately a自己 r fixation by formalin ment; penultimate segment with subdistal nar- solution. However,dark to light patterns in row orange band. Antenna1 p巴duncles each with coloration are generally well preserved even in fifth segment whitish; fourth segment whitish, formalin solution,in particular,in specimens with distal 0.3 -0.5 orange; third segment whit- just after the fixation. ish; second segment whitish,with distal 0.4 Distribution.- Kai Islands ,Indonesia; 0.5 orange except for white dorsomesial and ,Ryukyu ,and Amami Islands , dorso1ateral spines; first segment pale orange; Kagoshima,Japan. acicles with proximal 0.3 -0.5 orange,remain- der surfac 巴 whitish Right cheliped (Figs. 2D,3B) with both fingers generally white,each with subdistal nar- Turleania senticosa row transverse orange band and proximal dark (McLaughlin & Haig,1996) brown band. Palm generally faint orange with Figs. 2C-E ,3 , 7longitudinal orange stripes and i汀 egular distal orange area on dorsal face,very narrow dark Laurentia se nti cosa McLaughlin & Haig,1996: 87 , 白gs. 3E,6 brown areas on distal margins of dorsal and Turleania senti cosa- McLaughlin,1997: 477,figs. mesial faces,conspicuous dark brown patch 11 h-k. - Asakura,2004: 462,fig. 4. present at mesioproximal angle of dorsal face. ?Turleania similis Komai,1999: 50,figs. 27-31. Carpus generally faint orange;dorsal face with (See Remarks) ?Turl eania spinimanusKomai ,1999: 52 ,日gs. 32 faint orange transverse band medially,spines 35. - Komai etal. ,2002: 56. (See Remarks) on dorsomesial margin dark brown;mesial and lateral faces with few irregular faint orange Material examined.- 2 o ,SL = 1.9,2.3 areas. Merus general1 yfaint orange or whitish, mm,sledge net,off Kakeroma Island,Amami with few irregular faint orange patches on lat- Islands,st .11 ,RIV Toyoshio-maru,18 Nov. eral ,dorsal and mesiaI faces 1999,col l. Susumu Ohtsulくa,CBl¥ ιZ C 8781; Left che1iped (Figs. 2E,3 C) with both 日n- Io ,SL = 1.6 mm,200 m ,dredge ,off Onga- gers generally white,each with distal narrow zaki ,Ishigaki Island,Yaeyama Group,Ryukyu transverse brown band and proximal narrow Islands,RIV Yaeyama (cap.Koei Higa),21 dark brown band.Palm general1 yfaint orange, Nov.2002 ,col l. Takeharu Kosuge,CBM-ZC with 6longitudinal orange stripes and irregular 8782. Comparative specimens (for examination distal orange area on dorsal face,na 汀 owdark of morphological variation): 2o ,SL = 2.1 ,2.3 brown areas on distal margins of dorsal and me- mm,1 2 ,SL = 1.5 mm,Kai Islands,Indonesia , sial faces,conspicuous small dark brown patch KARUBARExpedition ,de t. P.A .McLaughlin , present at mesioproximaI angle of dorsal face. Pg 5270; 1o ,3 2 ,SL = 1. 6- 1. 8mm , Carpus general1 yfaint orange;dorsal face with Tanimbar Islands ,KARUBAR Expedition,de t. faint orange transverse band medially,spines P. A. McLaughlin,USNM 275999 ・2756000. on dorsomesial margin dark brown; mesial and lateral faces with few faint orange patches. D 目 cription 01c olor pattern (two days after Merus with some i汀 egular-shaped faint orange fixation by 10% formalin solution) .- Shie1d areas. Colorp att em so fhermit crabs 137

Fi g. 3. Turl eania sent icosa (M cL aughlin &H aig,1 996): ♂,SL = 1.6 mm,Is hi gaki-jim aI sland,Y aeyam a G roup,R yukyuI slands,C BM-ZC8782. A ,shi eld and cEphall ca ppendages (lER ante nnuleo rTImed),dorsa l; B ,r ightc helip ed,d orsa l;C ,left cheli ped,d orsa l;D ,le 代 second pereopod,la teral;E ,lef tt hird pereopod, latera l. Setae omittedin all ti gures .Sca les = 1mm . 138 A. Asakura

Second and third pereopods (Fig. 3D,E) oration from photograph of live animal and generally similar,each segment with transverse details from specimens two days after 白xation faint orange bands on semitransparent white in 10% formalin solution).- Shield (Fig. 4A) background.Dactyls each with subdistal ,me- mottled brown and white; ocular peduncles dial ,and proximal bands. Propodi each with each with darker area just beneath cornea;ocu- band in distal half and sometimes with irregular lar acicles whitish. patch on proximal half of lateral face. Carpi of Cephalic appendages (Fig .4A) gener- second each with medial and proximal bands; ally whitish,with orange transverse bands or 問中iofthird each with distal ,medial and prox- patches. Antennular peduncles each with distal , imal bands. Meri of second each with 2sub- medial,and proximal bands on ultimate seg- distal and 1proximal bands; meri of third each ment; penulitimate segment with median band. with band in distal half and in 什ont of proximal Antennal peduncles each with medial band on margm. fifth and fourth segments; third segment with Remarks.- See Asakura (2004) for mor- few small patches mesially and laterally ;first phological description of these specimens. 1 segment and proximal 0.2-0.3 of second seg- have no chance to examine coloration of live ment dark brown;antennal acicles with few animals. small brown patches dorsally The present study on morphological com- 80th chelipedswith chelae and 問中i bear- parison among the material examined (see ing longitudinal orange stripes on faint orange above) and similarity of coloration strongly background. Right cheliped (Fig.48 -0) with suggested that Turleania similis Komai,1999 , dactyl bearing 2stripes on dorsal face; very and Turleania spinimanus Komai,1999 ,could short stripe present ventroproximally on mesial be placed within morphological variation of face. Fixed finger with dorsal face bearing 2 T. senticosa .However ,it is not apurpose of stripes. Palm with dorsal face bearing 6stripes , the present study to evaluate specific status of 2fused into one and extending on fixed finge r. both Komai's taxa so that both species are here Carpus with dorsomesial margin bearing stripe , questionably placed under the synonymy of T. mesial face with 3stripes; lateral face with 2 senti cosa. stripes. Merus with medial transverse orange Distribution.- Seram,Kai and Tanibar band and several dorsal ,small iηegular-shaped Islands ,Indonesia; Yaeyama Islands ,Ryukyu , dark orange or brown areas . and Amami Islands,Kagoshima ,Japan . Left cheliped (Fig. 4E-G) with dactyl bear- ing 2stripes on dorsal face,mesial face with ventral stripe. Fixed finger with 2stripes on Catapagurus hirayamai (Asakura,200 1) dorsal face ,lateral one extending to dorsolat- Fig.4 eral face of palm. Palm with dorsal face bear- H emipagurus hirayamai Asakura,2001: 835 (key) , ing 6stripes including one from dorsolateral 879,table 1,figs. 15A-E,180 ,20E ,21 S-T,22J , margin of fixed finger.Carpus with stripe on 28E-H 30C 331 -L 35C 37C 39C 41 F-J 43E , , , , , , , dorsolateral margin,dorsomesial margin with F,441 ,51 .-Asakura,2002 ・450,451 (key) ,自 gs 1,20,3C. short stripe distally and proximally; lateral face Catapagurus hirayamai- McLaughlin,2004a: 13 . with 2stripes; mesial face with irregular orange areas .Merus with medial transverse orange Material examined.- 2 O ,SL = 2.0 ,2 .4 band and several dorsal small ,irregular-shaped mm,1 ov. 2 ,SL = 2.7 mm,off Shishihama, orange or brown areas. Numazu-shi,Shizuoka -ken, (= main Second (Fig. 4H,1) and third pereopods island of Japan),19 July 2003,coll. Show generally similar in coloration. Dactyls Hirayama,C8M ・ZC8783 with lateral and mesial faces each bearing longitudinal brown stripe on midline,lateral D escr卯 ion 01 color pattern (general col ・ face also with short brown line dorsally on 0.2 Colorpatt em so fhermit crabs 139

Fig. 4. Catapagurush irayamai (Asakura,2 001 ): o ,S L= 2. 0,Num azu- shi ,S hiz uoka-ken,C B M -ZC 8783. A ,shi eld and cephalic appendages,do rsa l;B ,rig ht chelip ed,la teral;C ,sa m e,do rsa l;0 ,sa m e,m es ial;E ,le 代 chelip ed,me sial;F ,sam e,d orsa l; G ,sa m e,la teral;H ,ri ght second pereopod,la teral ;E ,sa m e,m es ia 1. Setae omittedin a11 fi gures. Scales = 1mm . 140 A. Asakura

-0.3 from proximal end. Propodi each with ocular peduncles each with darker area just be- lateral face bearing 5 longitudinal brown neath comea and orange band anterior to proxi- stripes ,but ventral 2sometimes interrupted and mal margin; ocular acicles with orange areas on each separated into 2at 0.3 -0.4 from proximal lateral and mesial marginal areas. end; mesial face with 4 longitudinal brown Cephalic appendages (Fig.5A) gener- stripes ,but ventral 1or 2sometimes interrupted ally whitish,with orange transverse bands. and each separated into two; lateral and mesial Antennular peduncles each with band on 0.3 faces each with faint transverse orange band -0.4 from distal margin of ultimate segment medial1 yand proximal1 y; carpi each with lateral and additional band in front of proximal mar- face bearing broad,transverse orange band gin; penultimate segment with median band. medially,similar additional band proximally, Antennal peduncles with second and third seg- much darker dorsally and ventrally,and small ments much darker orange than other segments. dorsodistal darker area; mesial face with darker 80th chelipeds with chelae and carpi bear- area in proximal half,dorsoproximal portion ing longitudinal orange stripes on faint orange much darker. Meri each with lateral face background. Right cheliped (Fig. 5B-D) with bearing 4faint transverse orange bands,darker 2stripes on dorsal face of dacty l. Fixed finger dorsally and ventrally,and few additional dark orange.Palm and fixed finger with lateral small patches dorsally; mesial face with 3faint face bearing broad longitudinal orange stripe; and irregular transverse orange bands,darker dorsal face of palm with 6or 7stripes. Carpus dorsal1 yand ventrally,and few additional small with dorsomesial and dorsolateral margins each patches dorsally. with stripe ,lateral and mesial faces each with Remarks.ー-Coloration of live animal and additional stripe. 恥1erus with 2transverse or- that of specimens two days a仇er fixation in ange bands in distal half dorsally. forrnalin solution are very similar to each other. Left cheliped (Fig. 5E-G) with 2stripes on Distribution. -Westem Japan from Kyushu dorsal face of dacty l. Palm and fixed finger with to Honshu and Ogasawara Islands. dorsolateral margin bearing stripe ,palm with dorsal face bearing 3additional stripes and area between dorsolateral stripe and most lateral stripe with many short transverse lines. Carpus Catapagurus kosugei (Asakura,2001) with midline of dorsal face and dorsomesial Fig.5 and dorsolateral margins each with stripe; me- Hemipagurus kosugei Asakura,2001: 834 (key) , sial face with 1and lateral face with 2stripes. 854,table 1,figs. 9,170 ,190 ,21 G -H ,220 , Merus with irregular brown patches on dorsal 25E-L,29F -G ,32A -0 ,340 ,360 ,380 ,40L face . N ,42G -H ,440 ,51. - Asakura,2002: 450, 451 (key) ,白 gs .2A,3E. Second (Fig. 5H) and third pereopods gen- Catapagurus kosugei- McLaughlin,2004a: 13 . erally similar. Dactyls with dorsal and ventral margins orange. Propodi each with lateral face Material examined.- IO ,SL = 2.1mm , bearing 4 longitudinal orange stripes ,dorsal sandy mud bottom,13 - 16 m ,off mouth of one much darker; mesial face with no conspicu- Urauchi River,north west of Iriomote-jima ous color pattern .Carpi with brown stripes on Island ,Yaeyama Group,Ryukyu Islands,26 dorsal and ventral faces. Meri with lateral faces Aug. 2003,col l. Takeharu Kosuge,C8M-ZC bearing irregular pattems. 8784. Remarks.- Only preserved material was examined. 8ut,as suggested from the case of Description 01 color pattern (three days aι C. hirayamai,it would well represented color- ter fixation in 10% forrnalin solution).-Shield ation oflive animals. (Fig. 5A) mottled yellow-orange and white; Distribution.- Ryukyu lslands,Japan. Colorp a町 m s ofhennitc rabs 14 1

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Fig.5. Catapagurusk osugei (Asakura ,2 001 ): ♂,S L= 2. 1mm ,Iri omote-jim aI sland,Y aeyam aG roup, Ryukyu Islands,C BM-ZC878 4.A ,shi eld and cephali ca ppendages,do rsa l;B ,ri ght cheliped,la teral;C ,sa m e, dorsa l;0 ,sa m e,m es ial;E ,le ft chelip ed,me sial;F ,sa m e,la teral;G ,sa m e,d orsa l;H ,le ft second pereopod, late ra l. Setae omittedin a ll fi gures. Scales = 1mm . 142 A. Asakura

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Fig. 6. Nematopagurus gardineri Alcock,1905a: O ,SL = 2.80 mm,Numazu-shi ,Shizuoka-ken ,CBM -ZC 8785.A ,shie ld andcep hali cappendages ,dorsal; B ,left cheliped,lateral ;C ,same ,m esial; D ,sa me,dor sa l;E , left second pereopod,lateral; F,same ,mesial; G ,left third pereopod,lateral; H ,same ,me sia l. Setae omitted in al lfigure s. Scales = 1mm. Color patt ems ofhermit crab s 143

Nematopagurus gardineri Alcock,1905 lateral and mesial faces generally violet,paler Fig.6 distally and nearly white just beneath terminal claws,each with conspicuous subdistal nar- Nematopagurusgardi ne/'i Alcock 1905a: 834 p , , l. row pale brown band; mesial faces each with 68,fig. 3.- Alcock,1905b: 110,p l. 12 ,fig. 2 一一 McLaughlin,2004b: 212,fig. 23 or without very faint transverse bands medially Nematopaguru sholthui si McLaughlin and Hogarth, and proximally. Propodi whitish; lateral faces 1998 :25 ,白 gs. 19 -26. each withsingle brown longitudinal stripe and Nematopaguruspilosus Komai,1999: 28,figs. 14 - severalshort brown longit udinal stripes on 18. subdistal portion and on proximal half,these Not Nematopagurus gardineri一 Miyake,1978: 130 (i ncertae sedis ,see McLaughlin,2004b) sometimes fused,forming transverse bands; mesial faces of second each with 4or 5short Material examined.-lo ,SL = 2.80 mm, longitudinal stripes distally and proximally, sandy bottom,20 m ,off Shishihama,Numazu- mesial faces of third each with irregular brown shi ,Shizuoka-ken ,Honshu (= main island of areas distally and proximally. Carpi each with 5 Japan),20 Sept. 2003,col l. Show Hirayama, brown longitudinal broad stripes on lateral face ; CBM-ZC 8785. mesial face with some pale brown areas. Meri each with 4or 5broad and short longitudinal D esc/' iption 01 colo/' pattern (general col- stripes subdistally and,on lateral half,these oration from photograph of live animal and sometimes fused,forming transverse bands; details from specimen two days after fixation mesial face with some pale brown areas. in 10% formalin solution) .-Shield (Fig.6A) R emarks.- See McLaughlin (2004b) for mottled pale brown and white; ocular peduncles full synonymy. Coloration of live animaland each with broad deep brown band proximally that of specimens two days after fi. xation in for- with much darker area distally,whitish just malin solution are veηsimilar to each other. in front of band,gradually darker distally and Distribution .-Widely distributed in with very naηow deep brown band just beneath Indo-west Pacific :Cocos (Keeling) Islands, cornea; corneas silver with some brown spots; Maldives,Seychelles ,Kwa Zulu-Natal,South ocular acicles whitish. Africa,Japan ,New Caledonia. Antennules each with flagellum and uト timate segment violet; penultimate segment pale violet; basal segment pale brilliant red . A napagurus japonicus Ortmann,1892 Antennas each with ftagellum bearing alterna- Fig.7 tive white and brown bands; peduncle mottled Anapagu/'us pusillus var. japonica Ortmann,1 892: pale brown and white. Second and third maxil- 296,p l. 12 ,自 g.ll. lipeds with proximal portions brilliant red . Anapaguru sjaponicus 一 Miyake ,1961 :24. ー Color pattern of right and left chelipeds Miyake etal. ,1962: 126. 一 Miyake,1965 :648 , (Fig. 6B-0) generally similar. Chelae covered tig. 1099.-Miyake,1978 :139 ,text-tig .55 . Miyake& Imafuku,1980 :63 .- Miyake ,19 82: with dense gray setae. Both fi. ngers with distal 198 ,231. - Garcia-Gomez,1994 ・ 44,tig s.10 - halves white,each with subdistal narrow dark II .-Komai etal. ,2002: 48,fig. 12. brown band;single narrow longitudinal stripe present on mesial face of dactyl ,other po口ions Material examined.- 2 o ,SL = 1. 8,1 .8 pale orange,darker distally. Palm pale brown, mm,sandy bottom,20m ,off Shishihama, with some naηow dark brown areas on mesial Numazu-shi,Shizuoka-ken ,Honshu (=main face .Carpi pale brown,darker proximally. island of Japan),20 Sept. 2003,col l. Show Meri with naηow to broad longitudinal stripes Hirayama,CBM-ZC 8786 on dorsal ,lateral and mesial faces. Second (Fig. 6E,F) and third (Fig. 6G, Descr似 ion 01 color pattern (general col - H) pereopods generally simila r. Oactyls with oration from photograph of live animal and 144 A. Asakura

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Fig. 7. Anapagurusjaponicus Ortmann,1892: ♂, SL = 1.8 mm,Numazu-shi ,Shizuoka-ken ,CBM-ZC 8786 A ,right cheliped,mesial; B ,left cheliped,dorsal ;C ,left second pereopod,lateral; D ,same ,mesi al;E ,left third pereopod,latera l. Setae omitted in all 自gures. Scales = 1mm Color patterns ofhermit crab s 145 details from specimens two days after fixation IsIs ,27 0 05'N,142 0 13 'E ,col l. H. Tachikawa, in 10% formalin solution)- Shield generally 4May 1995,CBM-ZC 7368. Paratypes:1 O , whiti shwith some pale brown areas; ocular pe- SL = 2.4 mm,6 m ,Kaname-iwa ,Chichijima , duncles generally whitish; corneas pale brown Ogasawara Isls ,col l. H .Tachikawa ,25 Mar. with some brown spots. 1995,USNM 1014761 ;2 o ,SL = 2.4,1. 7 Antennules with flagella and ultimate and mm,1 = 1.6mm ,1 m ,B yoblトdani, penultimate segments semitransparent pale Chichijima,Ogasawara Isls ,col l. H .Tachikawa , brown or greenish brown. Antennas with flagel- 10 Mar.1995 ,CBM-ZC 7369;1 o ,SL = 2.5 la bearing alternative white and brown bands, mm,1 m ,Byobu-dani ,Chichijima ,Ogasawara pedunc1 es mottled pale brown and white. Isls ,col l. H .Tachikawa ,30 Mar. 1995,CBM- Second and third maxillipeds generally white, ZC 7370. Other material :2 o ,SL = 1. 9,1. 5 with brown articulations. mm,1 = 1. 7mm ,on coral head,15 m , Right cheliped (Fig. 7A) generally whitish, Shimamawari,off Wabuka,Kushimoto-cho , with both fingers each bearing subdistal naηow Higashimuro-gun,Wakayama-ken ,Honshu brown band;some irregular pale brown or or- (= main island of Japan),4 Nov .2003 ,col l. ange areas on each segment. Keiichi Nomura,CBM-ZC 8787. Left cheliped (Fig. 7B) with chela and car- pus generally whitish,with both fingers each D escription 01 color pattern (general color- with subdistal na汀 owbrown band; some irreg- ation from photographs of live Ogasawara spec- ular,pale brown areas and small ,darker brown imens and details from preserved Kushimoto幽 areas present on dorsal facesof palm,carpus cho specimens one month after collection ,fixed and merus. in 10% formalin for 10 days then transferred to Second (Fig. 7C,D) and third (Fig. 7E) 70% alcohol). - Shield and posterior carapace pereopods generally similar. Dactyls generally generallymottled dark brown or dark green brown,paler distally,each with narrow white and very pale brown or pale green,with brown band on proximal margin. Propodi whitish, gastric pits; ocular acicles each with proximal each with 2broad transverse brown bands,pal- transverse brown or dark green band. er mesially and area between bands sometimes Cephalic appendages generally with dark pale brown. Carpi each with broad transverse brown or dark green longitudinal stripes or brown band,darker dorsally and laterodistally. patches on whitish background. Antennules Meri each with 2brown patches dorsally and, with stripes on ultimate and penultimate seg- on second,ventrally ,and sometimes with 2 ments. Antennal peduncles with stripes on fifth transverse pale to dark brown bands laterally. segments and irregular patches on second to Remarks.一一 Coloration of live animal and fourth segments and acicles. that of specimens two days after fixation in for- Both chelipeds and second and third pereo- malin solution are very similar to each other. pods generally pale brown or pale green with Distribution .-Western Japan from Tokyo dark brown or dark green longitudinal stripes Bay to Kyushu. and patches. Right cheliped (Fig. 8A-C) with mesial face of dactyl bearing 2stripes ,connect- ed proximally by short band. Palm and fixed Boninpagurus acanthocheles finger with lateral margin bearing dark longitu- Asakura & Tachikawa,2004 dinal stripes ,palm with dorsal face bearing 4or Figs.8-9 5stripes on proximal half and proximal naηow transverse band but sometimes unclear;mesial Boninpagurusa canthochelesA sakura & Tachikawa, , 2004:158 ,白 gs .I-7 face with large patch. Carpus with dorsal face bearing 4or 5stripes of various widths;mesial Material examined. - Holotype: ♂,SL = face with dorsal stripe and ventral dark area . 2.6 mm,Byobu-dani ,Chichijima ,Ogasawara Merus with subdistal transverse band and ir- 14 6 A. Asakura

Fig. 8 目 B oninpagurus acanth ochelesAsa kura& Tachikaw a,2 004: o ,S L= 1. 9mm ,Ku shim oto-c ho, W akayam a-k en,C BM-ZC8787. A ,ri ght cheliped ,lat eral;B ,sa m e,d orsa l;C ,sa m e,m esia l;0 ,left cheliped , mesial;E ,sa m e,la teral; F,sa m e,d orsa l;G ,ri ght se cond pereopod,latera l;H ,sa m e,m es ial;1, ri ghtthird pereopod,la teral;J, sa me,m es ia l. Setae omitt edin a ll fi gur es. Scales = 1mm Color pattems ofhermit crabs 147

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Fi g. 9. Boninpagurus acanthochelesAsakura &Tachikawa ,2004: Kushimoto-cho,Wakayama-ken ,CBM-ZC 8787.A ,B ,D ,F ,G ,♂, SL = 1.9 mm;E ,o ,1.5 mm; C ,2 ,SL = 1.7 mm. A ,shield and cephalic appendages, dorsal; B ,third maxillipeds (meri through coxae) and anterior portion of sternite ,ventral ;C ,coxae and sternite of third pereopodsof female;D ,E ,coxae and stemite of fifth pereopods of male;F ,ri ght fourth pereopod,chela ,carpus and di stal portion of merus,lateral ;G ,telson ,dorsa l. Scales for A ,B ,D ,E = 1mm and for C ,F, G = 2mm. regular dark area on dorsal ,mesial and lateral 4stripes ,mesial faces of meri with few faint faces. patches. Left cheliped (Fig.8D-F) with dactyl bear- Remarks.- The males collected from ing 2stripes on dorsal face and 1additional Kushimoto-cho have ashort or very short ventral stripe on mesial face. Palm with dorsal , right sexual tube (Fig. 9D,E) like that of mesial and lateral faces bearing several irregu- Anapagrides de Saint LaurenトDechance,1966. lar stripes or pattems. Carpus with some stripes However,this species has multiple rows of on dorsal ,mesial and lateral faces. Merus with corneous scales in the propodal rasps of the irregular pattems on dorsal and mesial faces. fourth pereopods (Fig. 9F) (vs. single row of Second (Fig. 8G-H) and third (Fig.8I - J) scales in Anapagrides) and,a female speci- pereopods generally similar; lateral faces of men has paired gonopores (Fig. 9C) (v s. single dactyls,propodi ,and carpi each with 4stripes , left gonopore in Anapagrides). By overall very faint transverse,pale brown or green band morphological characters including these two also present on lateral faces of proximal por- and coloration,1 assigned these specimens to tions of dactyls,proximal halves of propodi , Boninpagurus acanthocheles. The right sexual distal portions and proximal halves of meri; tube of B. acanthocheles is medium len gth ,ca. mesial faces of dactyls,propodi ,and ca巾 ieach 2- 3coxal length,according to acriterion by with 3stripes; lateral faces of meri each with McLaughlin (2003),with atransparent distal 14 8 A. Asakura

po口ion .The sexual tubes of the present spec ト to Rafae lLem aitr e(NMNH) and N guyen menshav eno transparent portion ,but thi si s Ngoc-Ho (MNHN) for the opportunity to ex- most likely because the animals are not yet amine comparative m aterial duringmy stay in fully mature. As has been shown in Asakura th eir respective muse ums.Careful reviews by and Tachikawa (2004; Fig.6) ,the sex ual tube is Patsy A. McLaughlin (Shannon Poi nt Marine fully developed in the specimens over 2.36 mm Center,West ern W as hin gton University) , in SL,but the present specimens are smaller, Jose ph Poupin (Systとmes d 'lnformation SL= 1.5 ,1.9 mm. Geographiques In st itut de Recherche de l'Ecole Distributi on.- Japan: O gasawaraI slands Navale),and Stephen G. Dunbar (Department and now,Ku shomoto,Wakayama Prefecture . of Earthand 8iologic alSciences ,Loma Linda University) greatly improvedthe manuscrip t. Trichopagurus trichophthalmus (Forest,1954) LITERATURE CITED Catapaguroides? tri chophthalmus Forest ,1 954: 74, figs. 20-24. A lcock A. ,1905a. Marine cru staceans. XI Y. Trichopagurus tri chophth almus- de Saint Laurent, Pagurid ae. In :J. S. Gardiner,(e d. ),The Fauna 196 8: 927. - de Saint Laurent ,1970: 210 ,figs .1 and Geography of th eMaldives and Laccadive 16. - Asakura,2004: 460,tig. 3. Archipelagos,being anAcco unt of the Work carried out an dthe Coll ectio nm ade bya n Material exam ined. 一一 l ♂, SL=1 .4 0mm , Expediti ondurin gthe Years 1899 and 1900,2 : 827-835 ,p l. 68 3m ,under stone,reef edge,Nishihama ,Aka ・ 0 Alcock A. ,1905b .Ca talogue of the Indi anDecap od jima,Kerama Group,Ryukyu lslands ,26 1O' N , Crustacea in th eCo llection sof th eIndian 0 127 17' E,22 Apr.1994 ,col l. Keiichi Nomura, Museum.P art 1 1. Anomura. Fasc. 1,P agurid es. CBM-ZC 8788. Indi anMuseum ,Ca lcutt a. xi +1 97 pp. ,1 6pls . Asakura,A. ,2001. Arevision of the hemlit crabs of D escription 01 color pattern (白eld note by th ege neraCatapagurus A. Milne- Edwards an d Keiichi Nomura and specimen 6days after H emipagurus Smith fro m the Ind o- W est Pac i自c (C ru stacea :D eca poda :Anomura: Pagurida e) . fixation in 10% formalin). Chelipeds ambu- 一一 , Inv ertebrat eTaxonomy ,15 :823 -891 latory pereopods,antennas ,antennules ,ocular Asakura,A. ,2002. Rece nt topics on taxonom yof peduncles,and third maxillipeds totall yye llow her mit crabs from Japanese waters- family with no conspicuous stripe. Paguridae. Aquabiology (K aiyo tow Seibutsu , Seibutsu Kenkyusha) ,24: 449 56 R em arks.一一 See Asakura (2004) for mor- -4 Asakura,A. ,2004. Rece nt topi cs ontaxonomy of phological description of this specimen. her mit crabs from Jap anese waters- family Distribu ti on. - Tahiti ,Polynesia ;8ikini , Paguridae ,P ar tIY. Aquabiology (Kai yo tow Micronesia; Sulu,Sanguinishiapo ,lndonesia ; Se ibut su ,Seibutsu K enkyusha) ,26: 455-464 . RyukyuIslands ,Japan. Asakura,A. ,& Tachikawa,H .,2004. Boninpagurus 。canthocheles,anew genus and specie sof hermit crab (Decapoda: Anomura: Pagurid ae) Acknowledgements.- lgratefully acknowl- from shallow waters of the Ogasawara (Bonin) edge efforts by the followingpersons to collect Islands,J apan. Journ alof CrustaceanBiol ogy, specimens and take photographs,making this 24: 15 7-167. studypossible ;Takeharu Kosuge and Koei Higa Forest , }., 1954. C r凶 aces D ecapodes ma川 eur s shi gaki 8ranch Seikai National Fisheries des lI es deTahiti et desTu am otu .1 .Pa gurid ea (l , (suite). Bulletin du Museum National d'Hi stoire Research In stitute) ,Susumu Ohtsuka and staff Naturelle ,2e se ri e,26: 71-79. of R /V Toyoshio・maru (Takehara Station , Gracia-Gomez,J .,1994. The syste m atics of the Biosphere Science,Hiroshima U niversity) , genus A napagurus Henderso n,1 886,a nd a Show Hirayama (Numazu-shi),Hiro yuki new ge nus for Anapagurus drachi Fore st,1966 (C ru st ace a: D ecapoda :Pa gurid ae). Zoologische Tachikawa (Coastal Branch of Natural History Verhandelinge n,Leiden ,295 :1- 131 ,figs. 1-44. Museum and Institute ,Chiba) ,Keiichi Nomura Knowlton ,N. ,1 986.C ryptic and siblin gspecies (Kushimoto Marine Park). 1am also indebted am ongthe decapodCrustacea .J ourn alof Color patterns ofhermit crabs 149

Crustacean Biology,6: 356-363. Japanese) Knowlton,N. ,1993. Sibling species in the sea. Miyake,S. ,1965. Anomura. In: New Illustrated Annual Review of Ecology and Systematics,24 ・ Encyclopedia of the Fauna of Japan. 189-216 Hokuryukan,Tokyo ,(2) :630 -652,tigs. 1032 Komai,T .,1999. Hermit crabs of the families 1115. (I nJapanese) Diogenidae and Paguridae (Crustacea: Decapoda Miyake,S. ,1978 .The Crustacean Anomura Anomura) collected during the Shin'yo-maru of Sagami Bay. 200 pp. ,4 pls. Biological cruise to the Ogasawara Islands and Torishima Laboratory,Imperial Household,Tok yo Island ,Oceanic Island in Japan. Natural History Miyake,S. ,1982. Japanese Crustacean Decapods Research Speciallssue,6: 1-66. and Stomatopods in Color. 1. Macrura,Anomura Komai,T. ,Ohtsuka ,S. ,Nakaguchi ,K. ,& Go,A. , and Stomatopoda. 261 pp. Hoikusha Publishing 2002. Decapod crustaceans collected from the Company,Osaka. [In Japanese). southern pa吋 of the in 2000-2002 Miyake,S. & Imafuku,M. ,1980. Hermit crabs using TRV Toyoshio-maru. Natural History from the Kii Peninsula.Nanki-Seibutsu ,Nanki Research,7: 19-73. Biological Society,22: 1-7,59-64[In Japanese] . McLaughlin,P. A.,1997 .Crustacea Decapoda: Miyake,S .,Sakai ,K. ,& Nishikawa,S. ,1962 .A Hermit crabs of the family Paguridae from the fauna-list of the decapod Crustacea from the KARUBAR cruise in Indonesia. In: A. Crosnier coasts washed by the Tsushima warm current. and P.Bouchet ,(eds.) ,Resultats des Campagnes Records of Oceanographic Works in Japan, MUSORSTOM,volume 16,Memoires du Special Number,6: 121 - 13 1. Museum National d'Histoire Naturelle (Paris) , Ortmann,A. ,1892. Die Decapoden-Krebse 172: 433- 572. des Strassburger Museum,mit besonderer McLaughlin,P. A.,2003. IlI ustrated keys to families Beruksichtigung der von Herm Dr.Doderlein bei and genera of the superfamily Paguroidea Japan und bei den Liu-Kiu-Inseln gesammelten (Crustacea: Decapoda:Anomura) ,with diagnoses und zur Zeit im Strassburger Muse um of genera of Paguridae.Memoirs of Museum aufbewahrten Formen. IY. Die Abtheilungen Victoria,60:111 - 144. Galatheidea und Paguridea. Zoologische McLaughlin,P. A.,2004a. A reappraisal of the Jahrbucher,Abteilung ぬr Systematik,6: 241 hermit crab genera Catapagurus A .Milne- 326,pls. 11 - 12. Edwards and Hemipagurus Smith (Crustacea Rahayu,D. L. ,& Forest,J .,1999. Sur le statut Decapoda: Anomura: Paguridae),with the de Calcinus gaimardii H. Milne Edwards description of anew species. Zootaxa,433: 1- (Decapoda,Anomura ,Diogenidae) et description 16. de deux especes nouvelles.Zoosystema ,21: 463 McLaughlin,P. A.,2004b. A review of the hermit -472. crab genus Nematopagurus A .Milne-Edwards Saint Laurent,M. de.,1968. Revision des genres and Bouvier,1892 and the descriptions of tive Calapaguroides et Cestopagurus et description new species (Crustacea: Decapoda: Paguridae). de quatre genres nouveaux. 1. Catapaguroides A. In: B. Marshall & R. de Forges (eds.) ,Tropical Milne Edwards et Bouvier et Decaphyllus nov Sea Benthos,volume 23,Memoires du Museum gen. (Crustaces Decapodes Paguridae). Bulletin e National d'Histoire Naturelle (Paris) ,172 ・151 - du Museum National d'Histoire Naturelle,2 229. serie ,39 ['‘ 1967"]: 923-954,figs. 1-32 McLaughlin,P. A.,& Haig,J .,1996 .A new genus Saint Laurent,M. de. ,1970. Revision des genres for Anapagrides De Saint Lauret-Dechance, Catapaguroides et Cestopagurus et description 1966 (Decapoda: Anomura: Paguridae) and de quatre genres nouveaux.V. Trichopagurus de descrpition sof four new species. Proceedings of Saint Laurent (Crustaces Decapodes Paguridae). the Biological Society of Washington,109: 75- V I. Conclusion. Bulletin du Museum National d' 90. Histoire Naturelle,2e serie ,42 :210-222. McLaughlin,P .A. ,& Hogarth,P. J. ,1998 .Hermit Saint Laurent-Dechance,M. de. ,1966. Remarques crabs (Decapoda: Anomura:Pa guridea) from the sur la classi 自cation de la famille desPaguridae Seychelles. Zoologische Verhandelingen,Leiden , et sur la position systematique d'Iridopa gurus 318:1 -48 de Saint Lauren t. Diagnose d'Anapagrides gen. Milne Edwards,H .,1848 .Note sur quelques nov. Bulletin du Museum national d'Histoire nouvelles especes du genre Pagure.Annales des Naturelle,2< se rie ,38: 257-265. Sciences naturelles ,Pari s(Zoologie ,3 e Serie) , 10: 59 4 -6 Address:Natural History Museum and Institute , Miyake,S. ,1961. Decapod Crustacea. Fauna and Chiba,955-2 ,Aoba-cho ,Chuo ・ku,Chiba 260-8682 Flora of the Sea around the Amalくusa Marine Japan;e-mail: asakura@ chiba-muse.or. jp Biological Laboratory,2: I- iv ,1 - 30. (In