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Pomatocheles Je併の， Sii Miers，1879: the First Information on Pylochelid Larva (Anomura: Pylochelidae)

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Pomatocheles Je併の， Sii Miers，1879: the First Information on Pylochelid Larva (Anomura: Pylochelidae) CRUSTACEAN RESEARCH，NO .29 :65-69 ，2000 Hatchling of the symmetrical hermit crab Pomatocheles je併の， sii Miers，1879: the first information on pylochelid larva (Anomura: Pylochelidae) Kooichi Konishi and Michio Imafuku Abstrαct.-The hatchling larval stage Thus some previous authors regarded of the symmetrical and primitive hermit this family as the most primitive group crabPomαtochelesje伊 'eysii Miers，1879 among the hermit crabs (e.g. Barnes， was obtained under laboratory condi- 1980). Forest (1987) listed 40 species tions. This is the first recorded worldwide belonging to the Pylochelidae pylochelid larva. It has amass of yolk and described two post-larval specimens granules and the body is covered with a ofPomαtocheles stridulαns Forest，but no thin embryonic cuticle，showing a larval stages have been described to date. prezoea・like appearance. The antennal lndeed the Pylochelidae is the last ano・ exopod and mouthparts are reduced， muran family of which the larval stages while pereiopods，pleopods and the uro・ are undocumented，in spite of its signifi- pod are already well developed at the cance in terms of systematics. time of hatching. These features strongly In this paper we describe the hatching suggest that the larval development of P. larval stage of Pomatocheles jeffreysii je伊 eysii is abbreviated. Miers，and thereby provide the first infor- mation on pylochelid larvae. Introduction In the infraorder Anomura，phyloge- Material and Methods netic lineages in different taxa have been Wecollected an ovigerous female of widely discussed (e.g. ，McLaughlin ， Pomatocheles je庁'r eysii Miers by commer- 1983).Comparison of larval morphology cial trawling from the sea floor at adepth is one ofthe effective methods of phyloge- of 126 m off Anori，Mie Prefecture netic analysis as has been shown in the (45 0 12'N，135 0 40'E). All specimens of P. case of paguroids (MacDonald et al .， jefferysii inhabited tusk shells. The fe- 1957). Since Rathke (1840) confirmed the male carried only 5eggs ，with amean di- existence of metamorphosis in the life his- ameter of 1.4 3x 1. 11 mm，and was kept in tory ofhermit crabs，many carcinologists an aquarium at 10-120 C .One newly- have continued to describe anomur出 llar- hatched larva，slightly damaged in the val stages over the past one and ahalf- anterior portion，was fixed and preserved centuries.Their efforts have revealed the in 5% buffered formalin. The appendages features of the larval phase in all were dissected with fine insect pins or anomuran families except for the sym- sharpened tungsten needles under a metrical hermit crab family Pylochelidae Nikon SMZ-I0 stereomicroscope and (formerly Pomatochelidae). Unlike com- mounted on asilicon 幽 coated glass slide . mon hermit crabs，the general body plan Drawings and measurements were made of the pylochelids is s戸nmetrical and they with a drawing tube attached to an have segmented and calcified abdomens. Olympus BH-2 microscope.Carapace 66 K. KONISHI &M .IMAFUKU Fig.l. Pomαtochelesjeffreysii Miers，living specimens.A: adult removed 仕omthe host tusk shell ， B: larval stage just after hatching in dorsal view.An: antennae，e: compound eye. Scales 10 .0mm for Aand 1. 0mm for B . length (CL) was measured from the ante- Mandibles:Bulbous ，not clearly sepa- rior margin of the cephalothorax to the rated from the labrum and epistome. medial posterior border. Allillustrations Maxillule (Fig. 2E): Coxal and basial were made with Illustrator™ 5.5J (Adobe endites unilobed.Endopod unsegmented. 8ystems Inc.)using Macintosh™ 08 Maxilla (Fig.2F): Coxal and basial (Apple Co.Ltd. )computer . endites bilobed without setae. Endopod slender and unsegmented. 8caphogna- Results thite with several invaginated setae on its Ofthe 5developing embryos on the fe- anterior and posterior ends. male pleopods，only one hatched.The Maxilliped 1( Fig.2G ): Coxa and basis colour of the newly-hatched larva was red well developed，but with no setal arma- in the case ofliving specimen (Fig.1). The tures. Endopod 紅 ld exopod unsegmented， body and all of its appendages were cov- without setae. ered with thin embryonic cuticle ，and all Maxilliped 2(Fig. 2H):Endopod and setae were invaginated or folded within it ， exopod approximately equal in length. showing a “prezoea-like" appearance. Endopod incompletely 3-segmented with Carapace (Fig.2A ，B) :CL = 1. 31 mm. several invaginated setaedistally .Exo- Globular，without spines，and with a pod 2-segmented with 2terminal invagi- large mass of yolk granules beneath the nated setae. cephalothorax.Eyes large，stalked . Maxilliped 3(Fig .2 1): Endopod about Antennule (Fig. 2C): Endopodal ramus half length of exopod. about half length of exopodal ramus.Al l Pereiopods (Fig.2J-L) :Well developed aesthetascs and setae invaginated. in size ，pereiopod 1chelate ，and pereiopod Antenna (Fig.2D) :Endopodal ramus 4and 5smaller than the others. longer than exopodal ramus. Exopod re- Abdomen (Fig.2A ，B) :Six somites plus duced，covered with embryonic cuticle telson，somite 2-5 carrying buds of pleo ・ carrying one naked and 7setose processes pods. Somite 6with biramous uropods (2D，arrow). posteriorly. HATCHLING OFPOMATOCHELES JEFFREYSIl 67 E H Lur M' u (0 .1mm) C ，D，G ・| ーーーー E. F (O .5mm) J-L (1 .0mm) A ，B M Fig.2 .Pom αtocheles je庁， 'r eysii Miers，first larval stage just a氏er hatching: A ，whole larva in dor- sal view: B ，whole larva in lateral view; C ，antennule; D ，antenna ，aηow indicating embryonic culticle OndopodaI ramus;E，maxillule;F，maxilla;G，maxilliped 1;H;maxilliped 2;I，maxil- liped3iJ，pereiopod1(cheliped);K，pereiopod2;L ，pereiopod5;M，telson;M'，second processof telson.The areas hatched with lines in Aand Bindicate amass ofyolk granules. 68 K. KONISHI &M. lMAFUKU Telson (Fig. 2M): With small median length. Onthe other hand，the cuticle cov- cleft and 13 pairs of posterior processes. ering the antennal exopod shows an an- AlIprocesses including second (Fig. 2M')， tennal exopod (= scaphocerite) common to with marginal seta-like subprocesses. No anomuran prezoeas. It is concluded that 紅l Oillur 田 1 hair found. the prezoea-like larva of P. je庁'r eysii is considerably different from typical Discussion anomuran prezoeas. The presence of a The morphological features of the large amount of yolk granules and re- duced mouthparts suggest lecithotrophic newly-hatched larva of P. je所eysii found under the embryonic cuticle ，e.g. large development in this species. These fea- tures of P .je sii show that develop- body size 田 ld well-developed appendages かり similar to adult phase，strongly suggest ment after the first moult is atypical ab- that apost-larval form，megalopa or a breviated type. more advanced stage，would be released The reason for the lack of knowledge by the next moult. Setal invagination of concerning the larval forms of the Pylo- the cephalic appendages suggest that the chelidae may be the difficulty in obtaining present larva shows premature hatching. ovigerous females. Miyake (1998) listed 5 It may be，therefore ，inadequate to call pylochelid species belonging to five differ ・ this larva “prezoea" in the strict sense，i.e. entgenera 企omJapan. Most ofthem have the stage preceding the true first zoeal benthic habitats in mesopelagic depths stage. Gurney (1942) defined the term (Forest，1987; Yokoya，1933) ，and their prezoea for the larval stage when first reproductive cycle has not yet been stud- hatched and still covered by the embry- ied. Ecological studies from this point of onic cuticle .We hereafter use the term view are needed. Otherwise，for investiga- prezoea in the wide sense of Gurney's tion of the systematics of the pylochelid definition.Such a “prezoea stage" in ab- lineages，molecular studies would be also breviated development was observed by one of the most effective methods as Wear (1976) in Metanephros challengeri shown in the case of lithodid crabs (Balss)and some other decapod species. (Cunningham et al .，1992). In almost all decapods，this transient The present juvenile-like larval form last-embryonic stage is thought to be es- ofP.je伊， eysii itselfmay contribute alittle sential to the behavioural strategy adap- to our understanding of comparative lar- tive to hatching (Gore，1968; Konishi & val morphology in the Anomura，but we Quintana，1987). hope that typical larval forms wilI be Classification on the basis of prezoeal found in other species of this family. morphology in relation to adult taxonomy was first established by Lebour (1928) Acknowledgements based on antennal and telson processes in The authors are grateful to Takamichi brachyuran prezoeas. Hong (1988) Ando，Takeshima Aquarium (Gamagori， pointed out that anomuran prezoeas are Aichi Prefecture) for his kind help in ob- easily distinguished from other major de- taining the adult hermit crab. Our cordial capod groups by the outermost first and thanks are also extended Dr.P. A .Mc- second telson processes that were usually Laughlin for her valuable suggestions for naked and reduced to avarying extent. improving of the manuscript.We are However，this criterion cannot be applied deeply indebted to the late Dr. Sadayoshi to the present larva，because all the telson Miyake for his encouragement of our processes are setose and nearly equal in study. HATCHLING OFPOMATOCHELES JEFFREYSIl 69 Literature Cited Lithodes (Crustacea，Decapoda). Proceed- ings of the Zoological Society of London， Barnes，R.D. 1980. Invertebrate Zoology，4th 128: 209-257. edition. Saunders College Publishing， McLaughlin，P. A.， 1983. Hermit crabs - Are Philadelphia，1089 pp. they really polyphyletic? Journal of Crus- Cunningham，C.W .，Blackstone ，N.W. &Buss ， tacean Biology，3: 60ι621. L .W .，1992. Evolution of king crabs from Miyake，S. ，1998. Japanese crustacean deca・ hermit crab ancestors. Nature，355 ・539- pods and stomatopods in color. Vol. 1. 542. Macrura，Anomura and Stomatopoda. Forest，J. ，1987. Les Pylochelidae ou “Pagures Hoikusha，Osaka ，3rd printing，261 pp. 56 symetriques" (Crustacea Coenobitoidea). pls. [In Japanesel Memoires du Museum National d'Histoire Rathke，H.
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