Zoeal Development of Heteropilumnus Ciliatus (Decapoda: Pilumnidae), with a Key to the Known Pilumnid Zoeas in Korea and Adjacent Waters

JOURNAL OF CRUSTACEAN BIOLOGY, 23(2): 341–351, 2003

ZOEAL DEVELOPMENT OF HETEROPILUMNUS CILIATUS (DECAPODA: PILUMNIDAE), WITH A KEY TO THE KNOWN PILUMNID ZOEAS IN KOREA AND ADJACENT WATERS

Hyun Sook Ko and Hoi Jeong Yang

Department of Life Sciences, Silla University, Busan 617-736, Republic of Korea (corresponding author (HSK) [email protected]) Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021 ABSTRACT An ovigerous crab of Heteropilumnus ciliatus (Stimpson) was collected in South Korean waters, and the larvae were reared in the laboratory. Three zoeal stages are described and illustrated in detail. Morphological characteristics of the first zoea of H. ciliatus differ from those described by Takeda and Miyake (1968). The first zoea of H. ciliatus appears to be similar to the first zoeas of Heteropanope glabra Stimpson and Pilumnopeus makiana (Rathbun), but they can be distinguished from each other by differences of the carapace spines. A provisional key is provided to aid the identification of the pilumnid zoeas in Korea and adjacent waters.

In the family Pilumnidae, the complete larval drawings were made with the aid of camera lucida. Setal development has been described for eleven counts and measurements were based on about 10 specimens for each zoeal stage. The sequence of the zoeal description is species, whereas only the zoeal stages have based on the malacostracan somite plan and described from been described for five species. Ten of the anterior to posterior. Setal armature of appendages is de- sixteen species have been recorded in Korean scribed from proximal to distal segments and in order of and adjacent waters including Japan and China endopod to exopod (see Clark et al., 1998). The zoeal stages (see Table 1). are described and fully illustrated. For the second and third zoeal stages, only the main differences from the previous The pilumnid crab Heteropilumnus ciliatus stage are described. The long, plumose natatory setae of the (Stimpson, 1858) is found under stones or in first and second maxillipeds and the long antennular crevices of rocks in shallow waters (Sakai, aesthetascs were drawn truncated. The zoeal series and the 1976). The species is the only representative of spent female are deposited at Silla University, Korea. the genus found in Korea, and it also occurs on RESULTS the coasts of North China and Japan (Kim, 1973; Sakai, 1976; Dai and Yang, 1991). The Three zoeal stages occurred before meta- first zoea of H. ciliatus was first described by morphosis to the megalop stage. The durations Takeda and Miyake (1968), but that report was of the zoeal stages I–III at 258C were 4, 6, and 5 limited to brief comments and illustrations. The days, respectively. aim of this paper is to describe the zoeal stages Heteropilumnus ciliatus (Stimpson, 1858) of H. ciliatus in detail, to compare its morphology with previously described pilumnid Heteropilumnus ciliatus Takeda and Miyake, 1968: 127, fig. 3–4a–k, zoea I. zoeas, and to provide a key to the known pilummid zoeas in Korea and adjacent waters. Zoea I Figs. 1, 2 MATERIALS AND METHODS Size.—Carapace length 0.61 6 0.06 mm. One ovigerous crab of Heteropilumnus ciliatus was Distance from tip of dorsal spine to tip of collected by Hyun Sook Ko by SCUBA diving from Chuja Island, off the southern part of Korea, 2 July 2001 and rostral spine 1.93 6 0.05 mm. transported to a constant-temperature chamber at Silla University, Busan. The first stage zoeas hatched on 22 Aug Carapace (Fig. 1A, B, C).—Dorsal spine long, 2001 and were reared using methods described by Ko (1995) distally slightly curved and spinulate, longer at a constant water temperature of 258C. Larvae were fixed than rostral spine; rostral spine straight and and preserved in 10% neutral Formalin. Zoeal specimens slightly longer than antennal protopod; lateral were dissected using Leitz zoom stereomicroscope, and appendages were examined under a Leitz Laborlux S spines present and short; anterodorsal setae microscope. Appendages were mounted in polyvinyl lacto- absent; 1 pair of posterodorsal setae present; phenol, cover slips were sealed with clear nail varnish, and ventral margins without setae; eyes sessile.

341 342 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 23, NO. 2, 2003

Table 1. Earlier descriptions of larval development of species in the family Pilumnidae.

Recorded in Korea and Species Complete larval development Zoeal stages only adjacent waters Heteropanope glabra Stimpson Lim et al. (1984) Greenwood X and Fielder (1984a) P. lumpinus Bennett Wear (1967) P. novaezealandiae Filhol Wear (1967) Pilumnus dasypodus Kingsley Bookhout and Costlow (1979) P. vespertilio (Fabricius) Lim and Tan (1981) X P. hirtellus (Linnaeus) Salman (1982) P. minutus De Haan Ko (1994b) X

Parapilumnus trispinosus Sakai Ko (1994a) X Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021 B. eucratoides (Stimpson) (as Pilumnopeus eucratoides) Lim et al. (1986) X Benthopanope indica (de Man) Ko (1995) Wear (1968) X P. serratifrons (Kinahan) Greenwood and Fielder (1984b) Pilumnopeus makiana (Rathbun) Lee (1993) X Pilumnopeus granulata Balss Ko (1997) X Heteropilumnus ciliatus (Stimpson) Takeda and Miyake (1968) X Actumnus setifer (De Haan) Aikawa (1937) X Tanaocheles bidentata (Nobili) Ng and Clark (2000)

Antennule (Fig. 1D).—Uniramous; endopod Second Maxilliped (Fig. 2B).—Coxa without absent; exopod unsegmented, with 3 long (2 seta; basis with 4 setae; endopod 3-segmented, stout 1 thinner) aesthetascs, 1 short, slender with 1, 1, 6 (3 subterminal 1 3 terminal) setae aesthetasc, and 2 small setae, all terminal. respectively; exopod 2-segmented, distal segment with 4 terminal natatory plumose setae. Antenna (Fig. 1E).—Protopod slightly shorter than rostral spine and distally spinulate; exopod Third Maxilliped.—Present but small. slightly longer than protopod and distally Pereiopods (Fig. 2C).—Developing; cheliped spinulate on both lateral margins, with 1 larger bilobed. and 1 small medial spines. Endopod developing. Abdomen (Fig. 2E).—5 somites; somite 2 with Mandibles (Fig. 1F).—Asymmetrical; right pair of lateral processes directed laterally; molar with 4 teeth, left molar with 1 tooth, somite 3 with pair of lateral processes directed conﬂuent with incisor process; endopod palp posteriorly; somites 2–5 each with 1 pair of absent. posterodorsal setae; pleopods absent. Maxillule (Fig. 1G).—Epipod seta absent, Telson (Fig. 2D, E).—Each fork long and coxal endite with 7 terminal setae; basial endite spinulate, with 1 stout lateral spine, 1 minute with 5 setal processes; endopod 2-segmented, lateral seta and 1 dorsomedial spine; posterior proximal segment with 1 seta, distal segment margin with 3 pairs of stout spinulate setae. with 6 (2 subterminal 1 4 terminal) setae; exopod seta absent. Zoea II Figs. 3, 4 Maxilla (Fig. 1H).—Coxal endite bilobed, with 614 setae; basial endite bilobed, with 514 Size.—Carapace length 0.86 6 0.08 mm. setae; endopod bilobed, with 315(2sub- Distance from tip of dorsal spine to tip of terminal 1 3 terminal) setae; exopod (scaphog- rostral spine 2.18 6 0.08 mm. nathite) margin with 4 plumose setae plus distal Carapace (Fig. 3A, B, C).—2 pairs of ante- stout process. rodorsal setae present; each ventral margin with First Maxilliped (Fig. 2A).—Coxa without 5 plumose setae; eyes stalked; otherwise un- seta; basis with 10 setae arranged 2, 2, 3, 3; changed. endopod 5-segmented, with 3, 2, 1, 2, 5 (1 Antennule (Fig. 3D).—Exopod now with 6 long subterminal 1 4 terminal) setae respectively; (3 stout 1 3 thinner) terminal and 1 subterminal exopod 2-segmented, distal segment with 4 aesthetascs plus 1 terminal seta, otherwise un- terminal natatory plumose setae. changed. KO AND YANG: ZOEAL DEVELOPMENT OF HETEROPILUMNUS CILIATUS 343 Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021

Fig. 1. Heteropilumnus ciliatus, ﬁrst zoeal stage. A, lateral view; B, frontal view; C, lateral expansion of carapace; D. antennule; E, antenna; F, mandibles; G, maxillule; H, maxilla. Scale bars, A, B 5 0.5 mm; D, E, G, H 5 0.1 mm; C, F 5 0.25 mm. 344 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 23, NO. 2, 2003 Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021

Fig. 2. Heteropilumnus ciliatus, ﬁrst zoeal stage. A, ﬁrst maxilliped; B, second maxilliped; C, pereiopods; D, fork of telson; E, dorsal view of abdomen and telson. Scale bars, A, B, C, D 5 0.1 mm; E 5 0.5 mm. KO AND YANG: ZOEAL DEVELOPMENT OF HETEROPILUMNUS CILIATUS 345 Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021

Fig. 3. Heteropilumnus ciliatus, second zoeal stage. A, lateral view; B, frontal view; C, lateral expansion of carapace; D, antennule; E, antenna; F, mandibles; G, maxillule; H, maxilla. Scale bars, A, B, C 5 0.5 mm; F 5 0.25 mm; D, E, G, H 5 0.1 mm. 346 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 23, NO. 2, 2003

Antenna (Fig. 3E).—Endopod more developed; Maxilla (Fig. 5H).—Basial endite bilobed with otherwise unchanged. 616 setae; exopod (scaphognathite) margin with 19 plumose setae; otherwise unchanged. Mandibles (Fig. 3F).—Unchanged. First Maxilliped (Fig. 6A).—Coxa with epipod Maxillule (Fig. 3G).—Epipod seta now present; bud present; endopod terminal segment with 6 basial endite with 8 setal processes; exopod (2 subterminal 1 4 terminal) setae; exopod plumose seta now present; otherwise unchanged. distal segment now with 8 terminal natatory Maxilla (Fig. 3H).—Basial endite bilobed with plumose setae; otherwise unchanged. 615 setae; exopod (scaphognathite) margin Second Maxilliped (Fig. 6B).—Coxa with with 14 plumose setae, distal, stout process no epipod bud present; exopod distal segment Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021 longer prominent; otherwise unchanged. now with 8 terminal natatory plumose setae; First Maxilliped (Fig. 4A).—Exopod distal otherwise unchanged. segment now with 6 terminal natatory plumose Third Maxilliped (Fig. 6C).—Undifferentiated setae; otherwise unchanged. into segments; endopod longer than exopod; Second Maxilliped (Fig. 4B).—Exopod distal epipod present. segment now with 7 terminal natatory plumose Pereiopods (Fig. 6D).—Some segmental dif- setae; otherwise unchanged. ferentiation into segments. Third Maxilliped (Fig. 4C).—Biramous. Abdomen (Fig. 6G).—Somite 1 with 4 dorso- Pereiopods (Fig. 4D).—More developed. medial setae; somites 3–5 each with slightly longer posterolateral processes; pleopods Abdomen (Fig. 4F).—Now with 6 somites; (Fig. 6F) 1–4 with endopod buds; otherwise somite 1 with 2 dorsomedial setae; somites 2–6 unchanged. each with pleopods present, endopods absent; otherwise unchanged. Telson (Fig. 6E, G).—With pair of small dorsomedial setae; otherwise unchanged. Telson (Fig. 4E, F).—Posterior margin with 3 pairs of stout spinulate plus 1 pair of small setae. DISCUSSION Zoea III The description of the first zoea of Hetero- by Takeda and Miyake (1968) Figs. 5, 6 pilumnus ciliatus did not agree with many of the characteristics of Size.—Carapace length 1.01 6 0.06 mm. pilumnid zoeas given by Ko (1997). The first Distance from tip of dorsal spine to tip of zoea of H. ciliatus described in the present study rostral spine 2.60 6 0.05 mm. also differs significantly from that described by Takeda and Miyake (1968) (Table 2). Carapace (Fig. 5A, B, C).—Four pairs of Martin (1984) noted that the number of zoeal anterodorsal setae; each ventral margin with 7 stages of xanthoid species are 5, 4, 3, 2, and 0. or 8 plumose setae; otherwise unchanged. In the family Pilumnidae, the complete zoeal Antennule (Fig. 5D).—Biramous; endopod bud descriptions of 14 species are available; nine present; exopod with small proximal seta, 5 species (Heteropanope glabra, Pilumnus dasy- subterminal aesthetascs and 3 long and 1 short podus, P. hirtellus, P. minutus, P. novaezealan- slender aesthetascs plus 1 small seta, all diae, Parapilumnus trispinosus, Benthopanope terminal; otherwise unchanged. indica, Pilumnopeus granulata, and P. makiana) have four zoeal stages, and four species (Pilum- Antenna (Fig. 5E).—Endopodbudapproximate- nus vespertilio, Benthopanope eucratoides, ly 67 % length of exopod; otherwise unchanged. Heteropilumnus ciliatus,andPilumnopeus Mandibles (Fig. 5F).—Left molar with 2 teeth, serratifrons) have three. Pilumnus lumpinus has confluent with incisor process; endopod palp only one zoeal stage, and Aikawa (1937) de- present. scribed only the first zoea of Actumnus setifer, which already had buds of the third maxilliped Maxillule (Fig. 5G).—Coxal endite now with 9 and pereiopods. Therefore, it is not certain if this terminal setae; basial endite with 9 setal species has four zoeal stages or three zoeal stages. processes; otherwise unchanged. Some authors (Scotto, 1979; Rice, 1980) have KO AND YANG: ZOEAL DEVELOPMENT OF HETEROPILUMNUS CILIATUS 347 Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021

Fig. 4. Heteropilumnus ciliatus, second zoeal stage. A, ﬁrst maxilliped; B, second maxilliped; C. third maxilliped; D, pereiopods; E, fork of telson; F, dorsal view of abdomen and telson. Scale bars, A, B, C, E 5 0.1 mm; D 5 0.25 mm; F 5 0.5 mm. 348 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 23, NO. 2, 2003 Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021

Fig. 5. Heteropilumnus ciliatus, third zoeal stage. A, lateral view; B, frontal view; C, lateral expansion of carapace; D, antennule; E, antenna; F, mandibles; G, maxillule; H, maxilla. Scale bars, A, B 5 0.5 mm; C, F 5 0.25 mm; D, E, G, H 5 0.1 mm. KO AND YANG: ZOEAL DEVELOPMENT OF HETEROPILUMNUS CILIATUS 349 Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021

Fig. 6. Heteropilumnus ciliatus, third zoeal stage. A, ﬁrst maxilliped; B, second maxilliped; C. third maxilliped; D, pereiopods; E, fork of telson; F, pleopod; G, dorsal view of abdomen and telson. Scale bars, A, B, E, F 5 0.1 mm; C, D, G 5 0.5 mm. 350 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 23, NO. 2, 2003

Table 2. Comparison of the morphological characteristics of and the second maxilliped are unchanged during the first zoeas of Heteropilumnus ciliatus as given by zoeal development, it seems to be correct that Takeda and Miyake (1968) with those obtained in the present study. these two appendages should have eight and 1, 1, 6 setae, respectively. Hence, among the Takeda and pilumnid zoeas Heteropanope glabra, Hetero- Miyake (1968) Present study pilumnus ciliatus, and Pilumnopeus makiana Carapace closely resemble each other on the basis of Dorsal spine smooth spinulate zoeal morphology as follows: the carapace has Posterodorsal absent 1 pair short lateral spines and long dorsal and rostral setae spines; the endopod of maxillule and maxilla Antennule 2 aesthetascs 4 aesthetascs and and 1 seta 2 setae have 1, 214(6) and 8 setae, respectively; the Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021 Antennal 1 medial 2 medial spines endopod of second maxilliped has 1, 1, 6 setae; exopod spine and somites 2 and 3 of the abdomen have a pair Mandibles only left asymmetrical; right of lateral processes. The identification of the mandible molar with 4 teeth illustrated confluent with zoeas of the three species seems to be difficult incisor process when they are encountered as marine zooplank- Maxillule ton. We find that the characteristics of the Coxal endite no description 7 setae rostral and dorsal carapace spines are important to differentiate between these species. The zoea Maxilla of H. glabra is easily distinguished from those Basial endite 313 setae 514 setae Coxal endite 613 setae 614 setae of the other two species in having the rostral carapace spine about twice as long as the Maxilliped 2 antennal protopod, whereas in H. ciliatus and P. Basis 3 setae 4 setae Endopod 1, 1, 5 setae 1, 1, 6 setae makiana, the rostral carapace spine is equal in length or slightly longer than the antennal Abdomen protopod. Also, it is found that the only Somites 2–5 no description 1 pair of posterodorsal setae difference between the zoeas of H. ciliatus and Telson fork smooth, with spinulate, with 1 stout P. makiana is in the dorsal carapace spine, 1 lateral lateral spine, 1 minute which is spinulate or naked respectively. spine lateral seta, and 1 The following provisional key is for a rapid dorsomedial spine identification of the pilumnid zoeas in Korea and adjacent seas. The characteristics employed are consistent during zoeal development. suggested that the xanthoid species with less than four zoeal stages show abbreviated or advanced zoeal development and occupy somewhat re- Key to the Known Pilumnid Zoeas in Korea stricted estuarine habitats. The collecting site of and Adjacent Seas the present species, H. ciliatus, is in 8 meters depth at a small island far from the coast, and 1. Carapace with lateral carapace spines ...... 2 – Carapace without lateral carapace spines ...... 7 there is no low-salinity estuarine environment. 2. Rostral carapace spine about twice as long as antennal Therefore, we know only that pilumnid species protopod ...... Heteropanope glabra usually have four or sometimes three—rarely – Rostral carapace spine about equal or slightly longer only one—zoeal stages, and the reason for the than antennal protopod ...... 3 – Rostral carapace spine vestigial...... 4 occurrence of less than four zoeal stages is 3. Dorsal carapace spine spinulate...... unknown...... Heteropilumnus ciliatus The description of the first zoea of Pilumno- – Dorsal carapace spine naked. . . . Pilumnopeus makiana peus makiana by Lee (1993) is confusing. He 4. Lateral processes on somites 2, 3, 4, and 5 ...... described the first zoea as having the endopod ...... Pilumnus minutus – Lateral processes on somites 2 and 3...... 5 of the maxillule with seven setae and the 5. Endopod of second maxilliped with 1, 1, 6 setation. 6 endopodal setation of the second maxilliped as – Endopod of second maxilliped with 1, 1, 5 setation. 1, 1, 5, respectively. In the second zoea, the ...... Actumnus setifer former appendage was with eight setae, and in 6. Endopod of first maxilliped with 2, 2, 1, 2, 5 setation ...... Pilumnus vespertilio the third zoea, the setation of the latter – Endopod of first maxilliped with 3, 2, 1, 2, 5 setation appendage was 1, 1, 6. Considering that the ...... Pilumnopeus granulata characteristics of the endopods of the maxillule 7. Rostral carapace spine vestigial...... 8 KO AND YANG: ZOEAL DEVELOPMENT OF HETEROPILUMNUS CILIATUS 351

– Rostral carapace spine about equal or slightly shorter Lim, S. L., K. L. Ng, and W. H. Tan. 1984. The larval than antennal protopod length...... development of Heteropanope glabra Stimpson, 1858 ...... Benthopanope eucratoides (Decapoda, Xanthidae) in the laboratory.—Crustaceana 8. Lateral processes on somites 2, 3, 4, and 5 ...... 47: 1–16...... Parapilumnus trispinosus ———, ———, and ———. 1986. The complete larval – Lateral processes on somite 2 ....Benthopanope indica development of Pilumnopeus eucratoides Stimpson, 1858 (Decapoda, Brachyura, Pilumnidae) in the laboratory.— Crustaceana 50: 265–277. ACKNOWLEDGEMENTS ———, and W. H. Tan. 1981. Larval development of the hairy crab, Pilumnus vespertilio (Fabricius) (Brachyura, This work was supported by grant No. R04-2001-00022 Xanthidae) in the laboratory and comparisons with larvae from the Korea Science and Engineering Foundation. The of Pilumnus dasypodus Kingsley and Pilumnus sayi authors are grateful to Paul F. Clark (Natural History

Rathbun.—Crustaceana 41: 71–88. Downloaded from https://academic.oup.com/jcb/article/23/2/341/2679814 by guest on 24 September 2021 Museum, London) for his comments and criticism of a draft Martin, J. W. 1984. Notes and bibliography on the larvae of manuscript, and we thank Miss Se Jin Ok for help with drawings. xanthid crabs, with a key to the known xanthid zoeas on the western Atlantic and Gulf of Mexico.—Bulletin of Marine Science 34: 220–239. LITERATURE CITED Ng, P. K. L., and P. F. Clark. 2000. The Indo-Pacific Aikawa, H. 1937. Further notes on brachyuran larvae.— Pilumnidae XII, On the familial placement of Chlor- Records of Oceanographic Works of Japan 9: 87–162. odiella bidentata (Nobili, 1901) and Tanaocheles sten- Bookhout, C. G., and J. D. Costlow. 1979. Larval ochilis Kropp, 1984 using adult and larval characters with development of Pilumnus dasypodus and Pilumnus sayi the establishment of a new subfamily, Tanaochelinae reared in the laboratory (Decapoda, Brachyura, Xanthi- (Crustacea: Decapoda: Brachyura).—Journal of Natural dae).—Crustaceana, Supplement 5: 1–16. History 34: 207–245. Clark, P. F., D. K. Calazans, and G. W. Pohle. 1998. Rice, A. L. 1980. Crab zoeal morphology and its bearing on Accuracy and standardization of brachyuran larval the classification of the Brachyura.—Transactions of the descriptions.—Invertebrate Reproduction and Develop- Zoological Society of London 35: 271–424. ment 33: 127–144. Sakai, T. 1976. Crabs of Japan and the adjacent seas. Dai, A. Y., and S. L. Yang. 1991. Crabs of the China Seas. Kodansha Ltd. Japan. 773 pp., 251 pls. Springer-Verlag, Berlin, Heidelberg, New York, Tokyo. Salman, D. S. 1982. Larval development of crab Pilumnus 608 pp., 74 pls. hirtellus (L.) reared in the laboratory (Decapoda, Greenwood, J. G., and D. R. Fielder. 1984a. The complete Brachyura, Xanthidae).—Crustaceana 42: 113–126. larval development, under laboratory conditions, of Scotto, L. E. 1979. Larval development of the Cuban stone Heteropanope glabra Stimpson, 1858 (Brachyura: crab, Menippe nodifrons (Brachyura, Xanthidae) under Xanthidae), from Australia.—Australian Zoologist 21: laboratory conditions with notes on the status of 291–303. the family Menippidae.—Fishery Bulletin (U.S.) 77: ———, and ———. 1984b. The zoeal stages of Pilumno- 359–386. peus serratifrons (Kinahan, 1856) (Brachyura: Xanthidae) Stimpson, W. 1858. Prodromus descriptionis animalium reared under laboratory conditions.—Journal of Natural History 18: 31–40. evertebratorum, quae in Expeditione and Oceanum Kim, H. S. 1973. Anomura, Brachyura. Illustrated Encyclo- Pacificum Septentrionalem, a Republica Federata missa, pedia of Fauna and Flora of Korea.—The Ministry of Cadwaladaro Ringgold et Johanne Rodgers ducibus, Education, Korea 14: 1–506. observavit et descripsit W. Stimpson. Pars IV. Crustacea Ko, H. S. 1994a. Larval development of Parapilumnus Cancroidea et Corystoidea.—Proceedings of the trispinosus Sakai, 1965 (Crustacea, Brachyura, Xanthi- Academy of Natural Science, Philadelphia 10: 31–40. dae) reared in the laboratory.—Korean Journal of Takeda, M., and S. Miyake. 1968. First zoea of two Zoology 37: 331–342. pilumnid crabs of the family Xanthidae.—Science ———. 1994b. The zoeal stages of Pilumnus minutus De Bulletin, Faculty of Agriculture, Kyushu University 23: Haan, 1835 (Decapoda: Brachyura: Pilumnidae) in the 127–133. laboratory.—The Korean Journal of Systematic Zoology Wear, R. G. 1967. Life-history studies on New Zealand 10: 145–155. Brachyura 1. Embryonic and post-embryonic develop- ———. 1995. Larval development of Benthopanope indica ment of Pilumnus novaezealandiae Filhol, 1886, and of (De Man, 1887)(Decapoda: Brachyura: Pilumnidae) in the P. lumpinus Bennett, 1964 (Xanthidae, Pilumnidae).— laboratory.—Journal of Crustacean Biology 15: 280–290. New Zealand Journal of Marine and Freshwater Research ———. 1997. Larval development of Pilumnopeus 1: 482–535. granulata Balss, 1933 and Pilumnus minutus De Haan, Wear, R. G. 1968. Life-history studies on New Zealand 1835 (Crustacea: Brachyura: Pilumnidae), with a key to Brachyura 2. Family Xanthidae. Larvae of Heterozius the known pilumnid zoeae.—Korean Journal of Bi- rotundifrons A. Milne Edwards, 1867, Ozius truncatus ological Sciences 1: 31–42. H. Milne Edwards, 1834, and Heteropanope (Pilumno- Lee, D. H. 1993. Larval development of Macromedaeus peus) serratifrons (Kinahan, 1856).—New Zealand distinguendus (De Haan, 1835) and Pilumnopeus maki- Journal of Marine and Freshwater Research 2: 293–332. ana (Rathbun, 1929) (Brachyura, Xanthidae) reared in the laboratory.—Ph. Master Thesis, Busan National Univer- RECEIVED: 17 April 2002. sity, Busan, Korea. 53 pp. ACCEPTED: 23 September 2002.