(fa £i^x\' Cah. Riol. Mar. (1997) 38 : 29 I-299 v^C^I /
The first zoeal stage of Glyptograpsus impressus, with comments on the subfamilial arrangement of Grapsidae (Crustacea: Brachyura).
Jose A. CUESTA1 & Christoph D. SCHUBART 'Departamenta de Ecologfa, Facuitad de Biotogia, Lhuversidad de Sevilla, Apdo. 1095, F-4I0S0 Sevdlti/Spain Facidtdt fur Biologic I: VHF. Universitdt Bielefeld, Postfach 100131, D-33501 Bielefeld 7 Germany -' Present address : Department of Biology, University of Southwestern Louisiana Lafayette, LA 70504-2451/USA ! Fax 34 54 61 57 SO - E-mail [email protected]
Abstract: The first zoeal stage of the brackish-waier crab Glyptograpsus impressus is described and illustrated from laboratory hatched material. This is the first description of a larval stage from the Central American genus Glyptograpsus which is traditionally placed within the grapsid subfamily Varumnae. Comparison of zoeal setation patterns revealed common features with the Varuninae. while other features are more typical for the Grapsinae and Plagusiinae. Furthermore, zoeae of G. impressus are distinct in the setation of the maxillar endopod. which is unique within the Grapsidae and makes the inclusion of the genus Glyptograpsus within one of the presently recognized grapsid subfamilies questionable.
Resume : Le premier stade zoo du crabe d'eau saumatre Glyptograpsus impressus est deerit et illustre a partir de materiel eclos en laboratoire. Ceci represenle la premiere description d'un stade larvaire d'un genre de I'Amerique cenlrale. Glyptograpsus, qui est traditionnellement place dans la sous-famille des Varuninae. Une comparaison des soies dcs zoes revele des structures communes avee les Varuninae. tandis que d'autres caraeteres sont plus tvpiques des Grapsinae et des Plagusiinae. De plus les zoes de G. impressus sont differentes par les soies de I'endopodite maxillaire, unique parmi les Graspidae, ce qui remel en question la place du genre Glyptograpsus dans Tune des sous families de Grapsidae.
Keywords : larval morphology, zoea. systematies, phylogeny, er
Introduction describe this subfamily are ambiguously defined and it is widely believed that the Varuninae represent an artificial The brachyuran family Grapsidae is presently subdivided grouping (P. Davie, D. Guinot. M. Turkay, R. Manning into 4 subfamilies: Grapsinae Dana. 1851. Plagusiinae personal communications 1997). Dana. 1851. Sesarminae Dana. 1851 and Varuninae Alcock. The present taxonomy of the Brachyura is mostly based 1900. This taxonomic classification goes back to Alcock on adult morphological characters. Rice (1980) emphasized (1900) and has been adopted in subsequent systematic the relevance of larval morphology for phvlogenetie studies reviews of this family. The Varuninae comprise 22 within the Brachyura. which gained further evidence by morphologically and ecologically very diverse crab genera. recent studies {e.g. Clark & Webber. 1991; Marques & Several morphological characters used by Alcock (1900) to Pohle. 1995). In the Grapsidae, larval morphological characters have occasionally been related to their taxonomic Reai 1c 29 juillcl 1997: acceple apres revision le 27 jamier 1998. classification (Bourdillon-Casanova. 1960; Wear. 1970; Received 29 July 1997: accepted in revised form 27 January 1998. Wilson. 1980; Pereyra Lago. 1993) and allowed to show '»2 rikST/Ol-.AL SI'ACil-. Ol- CI ) I'lOCKM'SCS IMI'l-H-SSCS diagnostic characters for Ihc subfamih Grapsinae (Cuesta a from Ihc base to the tip of ihe lateral spines: antennal length dl.. 1997). Within the Sesarminae and Varuninae. most (al) from the base to Ihe lip of the spinous process. genera are also \er\ homogeneous in their larval Semipermanent mounts were made of whole larvae and morphology, while others show marked differences (Rice. dissected appendages were stained using CMC 10 and 1980: Percyra Lago. 1993; Sehubarl & Cuesta. in press). lignin pink. Due to the unusual setation pattern, more than Out of the 51 genera presently reeogni/ed within the 50 larvae were dissected for confirmation. OTIC female crab Grapsidae. larval stages have been examined only in 33 and a sample of larvae were deposited in the United States genera. The lack of knowledge applies particularly to the National Museum of Natural History under Ihe catalog Varuninae for which larval descriptions are only available number USNM cat. 2S4160. for 8 out of 22 genera. Thus, to elaborate a taxonomic svstem for the Grapsidae based on both adult and lar\al Results morphology, it would be necessary to obtain descriptions of Several observations on the reproductive biology of larvae from many more genera. Glxpt<>!>rapstts impressus were obtained during 18 months In this study, a larval stage of the grapsid genus of captivity. In 1996. prolonged copulations of the larger Glxpio^rapsus is described for die first time. This genus is male with the two females were observed without previous currently placed within the Varuninae and consists of two molls of the females [i.e. in hard condition) from July to species which are reslricled to Central America. The September. In July 1997. the surviving male and female morphology ol' the first zoeal stage of G. impressus Smith. remained in copula for two weeks. Females always became 1870 is compared with thai of olher Grapsidae. Based on ovigerous I to 3 days after copulations. They carried eggs these findings, the systematic position of the genus for 3 to 4 weeks. In 1996. the larger female of 20.6 mm Glxpio^rapsus is discussed. carapace width (cvv) copulated and carried eggs twice within the same molting cycle, while the smaller female Materials and methods (17.6 nun ew) only produced one brood. In 1997. the larger female (21.2 mm cvv) released larvae only once, before Adult (jlypi(>i>rapsns impressus were obtained from molting in September. This last hatch consisted of Fart an (Panama) in February 1996. without encountering approximately 3000 larvae. Fresh water was preferred for any ovigerous female in the field, and transported to ihe larval release in 2 out of 3 cases. In 1996. many larvae died University of Bielefeld (Germany). Two females and Hvo as pre/oeae. There were no significant differences in males were kepi in a 30 x 20 em aquarium with access to survival of first /oeal stage maintained in salinities of 16 or brackish water (1-5 PSU) and to rotting wood which was 32 PSU. Unfortunately no larva lived longer than eight days used as food and shelter. Temperatures ranged from 21 to and no successful moll to the second /oeal stage was 24 C and the light : dark cycle was 14 : 10 hours. In 1996, achieved. The small si/c of the larvae apparently did not females were placed in a separate aquarium following egg allow Ihem to feed on Anemia sp. nauplii. Detritus enriched extrusion and were offered small containers with water of 0 water or rotifer cultures (in 1997) also failed to extend and 16 PSU salinity for larval release. After hatching, survival. approximately 300 larvae were maintained in groups of 5 individuals and in mass-culture conditions (> 50 Description individuals). Light and temperature regimes were the same as described above. Differential survival was lesied in Glxpiotfrapsus impressus Smith. 1870 salinities of Id and 32 PSU. Samples of larvae were fixed in Zoea I (Figs. 1, 2) cither 5'/< formal in or HY/i ethanol. Dimensions: rd: 0.66 ± 0.03 mm: cl: 0.37 ±0.01 mm: cw: Drawings and measurements were based on 20 larvae 0.48+0.01 mm; ds: 0.21 ± 0.02 mm; rs: 0.16 ± 0.01 mm: and made using a Wild M/6 and Zeiss compound Is: 0.08 ±0.01 mm; al: 0.16 ±0.02 mm. microscope with Nomarski interference, both equipped with Carapace (Fig. 1A), Globose and smooth. Dorsal spine a camera lucida. All measurements were made through an slightly curved, thin rostral spine straight, both well ocular micrometer and refer to the following distances: developed. Lateral spines slightly curved and directed roslro-dorsal spine length 0x1): from the tip of the rostral anteriorly. Pair of dorso-lateral simple setae near base of spine to the lip of the dorsal spine: carapace length (cl>: dorsal spine. No selae on lateral margin. Eves sessile. from the base of Ihe rostrum to the posterior carapace Antennule (Fig. 1C). Uniramous. Endopod absent. margin; carapace width (cwl: distance between Ihe lips of Exopod unsegmented with two unevenly sized aesthetascs the lateral spines; dorsal spine length Ids): from the base to and two thin simple setae. the tip of the dorsal spine; rostral spine length (rs): from the Antenna (Fig. ID). Well developed protopod. almost base to the tip of the rostral spine; lateral spine length (Is) equal in length to rostral spine, proximally bearing two rows J. A. CUHSTA. C II. SCllliR \RT 24.'
Figure 1. ("ilvpio^rapsus impressits. /oca 1. A. lateral view: B. abdomen, dorsal view: C. anlennule: D. antenna. Scale bars = ().| mni. Figure 1. (jlvptograpsiis inipre.ssus, /oe I. A. vue laterale : B, abdomen, vue dorsale ; C. aiitennule : D, antcunc. Fxhelles = 0.1 mm. of denticles. Fndopod ahsent. Fxopod elongated about 3/4 Scaphognalhite with four plumose marginal setae and long the length of protopod. with three unequal simple setae and stout posterior process. one longer terminal seta. Hirst maxilliped (Fig. 2C). Basis with eight medial Maxillule (Fig. 2A). Coxal and basial endites each with plumodenticulate setae arranged 2.2.2.2. Fndopod 5- live plumodenticulate setae: no seta on external margin of segmented with 12 plumodenticulate or simple setae basis. Fndopod 2-segmented. proximal segment with one arranged 2.2.1,2,5. Fxopod slightlv constricted with four simple seta, distal segment with one subterminal and four long terminal plumose natatory setae. terminal plumodenticulate setae. Second maxilliped (Fig. 2D). Basis with four medial Maxilla (Fig. 2B). Coxal endite bilobed with 5+3 plumodenticulate setae arranged 1.1,1,1. Fndopod 3- plumodenticulate setae on the outer and inner lohe segmented with 8 plumodenticulate or simple setae respectively. Basial endite bilobed with 4+4 on the inner and arranged 1.1.6 (three subterminal and three terminal). outer lobe. Fndopod unsegmented, bilobed with one long Exopod slightly constricted with four long terminal plumose plumodenticulate seta on the inner lobe and two long setae natatory setae. (outer simple and inner plumodenticulate) on the outer lobe. Third maxilliped. Absent. :i)4 K1RST7QPAL S'I ACiK OF CLYPIOCRArM .S /.*//>A7/.S.S7'.V
Figure 2. Glxplo^rapsits impressiM, zoca I. A, maxiliule: B. maxilla: C. first maxilliped: D. second maxilliped. Scale bars = (Id mm. Figure 2. (ilvpto^rapsiis iiupressus. /oe I. A. maxiliule : B. maxillc : C, premier maxillipede : D. second maxillipede. Fxhelles = 0.1 mm.
Pereiopods. Absent. subfamilies (see Bourdillon-Casanova. I960; Wear. 1970; Abdomen (Fig. IB). Five abdominal somites; dorso Wilson, 1980: Rice, 1980). Table 1 summarizes data on first lateral knobs on somites 2-4 (smaller on somite 4). Somites zoeal features of all described Varuninae and Plagusiinae 2-5 with a pair of sparsely plumose setae on posterio-dorsal species (for comparison with Grapsinae see Cuesta et al. margin. Pleopods absent. (1997) and with Sesarminae see Pereyra Lago (1993)). The Tclson (Fig. IB). Six serrulate setae on posterior margin first zoea of G. iiupressus shows two characteristics (type B of antenna and telson) which are consistently found in the of telson. No lateral spines on outer side of furcae. Furcae Varuninae. The strongly developed antennal exopod is also covered with two rows of small setules. characteristic for the Varuninae (and Sesarminae). On the other hand. G. iiupressus larvae differ from the Varuninae in Discussion several of the setation patterns. The 2,2,2.2-setation on the First zoeal morphology of Gixpiograpsiis inipressiis basis of the first maxilliped is typical of the Plagusiinae and does not allow to place this species in one of the existing Grapsinae, while the 1.1.6-selalion on the second maxilliped .1. A. CL'ESTA. C. ft Snil'HART 295
Table 1. First /oeal lealnrcs of all known Varnninae, including (ilyptn»rnpsus impressu\ and Plngusiinae. Tableau I. Caracteres des premieres /oes de tons les Varunmae connus. \ coinpris (jl\pio»nipsit\ imprrwits et des Rlagusiinae.
No. of Telson Antenna Lateral Maxilla Maxilliped Maxiliiped 2 Abdominal Specie- Zoeal Carapace References Stages Type Type Spines Endopod Basis Endopod Endopod Knobs
Acmtteop/euni parvula 5(6} B B (3) 2.2.3.3 0.1.6 Kim & Jang, 19S7
Caieliee depressus 5 B B(3i 2,2,3,3 0.1.6 2-3 Kim At Lee. 1483 limehwouis titlanliciis 5 B B(3l 2.2,3.3 2.2.1.2.5 0.1.6 Rodriguez^/.. 1992 Brachynotus ^cmmclkiri 5 B B t.1) 1 ^ 3 3 2.2.1.2.5 0.1.6 Gnerao el ui.. 1995 Brat hynotus sexdcnlalus 5 B B(3) 2.2.3.3 2.2.1.2.5 0.1.6 Cnesla & Rodriguez, unpublished data
Cvrio^rapMis ttltinmnus 5 B B<3) 2.2.3.3 2.2.1.2.5 0.1.6 2-3 Scel/o Sc Lichlschem deBastida. 1979 C\IH>\>U\pMt\ url^altllllS 5 13 B (3) 2.2.3.3 2.2.1,2.5 0.1.6 Bosebi. 1981
F.riocheir lepu>v,miihu\ 5 B B(2) 2.2.3.3 2.2.1.2.5 0.1.6 "M Lee. 198K Friacheir sinensis 5 B B(3) 2 2 2.2,3.3 2.2.1.2.5 0.1.6 Kim& Hwang. 1995' F.riocheir japtmit us 5 B B(3) i i 2,2,3.3 2.2.1.2.5 0.1.6 2-4 Kim& Huang. 1990 •i triucheir jormosu 5 B BC) 1.1.1.1.4 5 2-4 Sin A: Yu. 1992
Hemi^rapsus nudtis 5 B B(3) 11 1.2.3.3 2,2,1.2.5 0.1.6 2-3 Hart. 1935 Heini^rapsiis ore^onensis 5 B B(3) 1.2.3.3 2.2.1.2,5 0,1.6 Hart. 1935 IIemi<>rapsns penicillatus 5 B B (3> 2.2.3.3 2.2.1.2.5 0.1,6 2-3 Terada. 1981: Hwang & Kim. 1995 Henti^rtipsus sanguineus 5 B BCD 2 2 1 ^ 3 3 2.2.1.2.5 0.1.6 2-3 Hwang etui. 1993 Hemigrupsus sinensis 5 B B(3i 2 2 2.2.3.3 2.2.1.2.5 0.1.6 2-4 Kim & Moon. 1987 Hcniiiimpstts longitursis 5 B B (3) -i -i 2,2.3.3 2.2.1.2.5 0.1.5 Terada. 1981 ficnii^rapsus crenulatus 5 B B(3) -, -i 3.3.3.3 1,1.1.2.5 1.1.6 Wear, 1970 Henii^rapsus etlwardsi 5 B B(3) -i i 3.3.3.3 2.2.2.2.6 1.1.6 Wear, 1970 11 ; ; B B(3) -> 7 2.2.1.2.5 0,1,6 Cuesta el til. (unpublished}
(ihproiirapsus impressus '.' B Bt4t 2,2.2.3 2.2.1.2.5 1. 1,6 2-4 present studs tucliiro^rapsus amerkanus 5(6) A- ('(2) 2 2 2 2 2.2.1.2.5 1.1.6 2-5 Wilson. 1980
Hagusiinae ASM29 '.' A C(2) 2.3 1 2 2 2 2.2.1.2.5 1.1.6 2-5 Rice& Williamson. 1977
Pltiifitsiti d. ih'presstt 67 B C(2) -> 3 -• i i i 2,2,1,2,5 1.1.6 2-5 Wilson
Peivnon i>ibl>esi i~> i T 2.2.1.2.5 Lebour, 1944: Paula & Hartnoll. 1989
Tabic 1. Telson and antenna type according lo Aikawa (1929}. In column "Antenna Type", numbers in brackets correspond to the number of setae on the exopod. In column "Lateral Carapace Spines'": + refers to presence. +! to presence with tubercles, spines and/or setae,- to absence. Signs '.' = no data. Exponent numbers ' to ••' mean: '. Larval development also described by Montii ft al. {1996) tor a German population of triocheir sinensis, introduced earh this century into Europe: several differences were found and here we prefer to use the description from the original Asian population; -. type B in subsequent /oeal stages: •', 1.1.5 in Aikawa's description: 4, 2-4 in Aikawa's description; -\ plankton-collected, instar number estimated Irom the sizes of the /oeae: (l. deduced from illustration, first group of setae not visible: 7. 2.3 in subsequent stages, with one plumose seta. Tableau 1. Type dc telson et d'antenne d'apres Aikawa (1929}. Dans la colonne "Type d'antenne". les nombres entre parentheses correspondent an nombre de soies sur I'exopodile. Dans la colonne "Fpines laterales de la Carapace": + sjgnifie presence, +! presence avec tubercules. epines et/ou soies, - signiiie absence. 1 - pas de donnees. Les exposants ' a 7 signifient : '. devcloppement larvairc egalemenl decrit par Montii el al. (1996). sur une population d'Allemagne. de triocheir .sinensis, espece introduite an debut du siecle en Europe : plusieurs differences out ete note'es el iei nous prelerons utiliser la description basee sur la population asiatique : -, type B dans les stades zoes suivants ; -\ I, 1.5 dans la description de Aikawa ;4, 2-4 dans la description de Aikawa. :\ stades /oes recol- tes dans le plancton. nombre de stades estimc d'apres la tailie des larves ;(>. deduit d'une illustration, premier groupe de soies non visible. •'. 2-3 dans les stades suivants. avec une soie plumeuse. :% FIRS'I /.OhAI.STACih OK CI.Yl'HHiKM'SCS IMI'KLSSCS endopod is only found in Pla^usiu, Euchiro^rapsiis and probably represents a zoea of Euchiroi>rapstis as suggested Ci!\ptoi>rapsus. Wear (1970) also described a 1.1.6-setation by the morphology of the telson (Table I: see also Paula. for two species of Hewi^rapsus from New Zealand. 1987). According to Rice (1980). the Plagusiinae larvae are H. edwardsi (Hilgendorf. 1882) and //. creiudciius characterized by: 1) a well-developed lateral spine which, (H. Milne-Edwards. 1837). However, recent re-examination like the rostral spine, carries tubercles, spinules or setae. 2) of H. edwardsi lirst zoea! stage revealed the more typical a short or rudimentary antenna! exopod. 3) 2.3-setalion on 0.1.6-pattcrn {Table 1). The second species of this genus the maxillary endopod. and 4) basal segment of the second described by Wear (19701. H. crenulatiis, is now die only maxilliped endopod armed with a single seta. Wilson (1980) Hemigrapsiis with the questionable 1,1,6-setalion and a re described small rounded protuberances on the dorsal examination ofzoeae of this species needs to be done. carapace posterior to the dorsal spine (see Fig. 3) as an The lirst /oeal stage of Glyptograpsus impressits has a additional common character to the Plagusiinae. This 1.2-setalion on the endopod of the maxilla which is unique character was also found in "Plagusiinae ASM29" - Rice among all the known zoea larvae of the Grapsidae. While a and Williamson. 1977 and Percnon s>ibbe.\i (Lebour. 1944), setation of 2.3 is common for Plagusiinae and Sesarminae. but was not mentioned in the descriptions of Aikawa (1937). Varuninae and Grapsinae have a 2.2-pattern (Rice. 1980). Wear (1970). Terada (1987) and Paula & Hartnoll (1989). The 1.2-setalion is otherwise onlv found in the families Instead, the last two authors proposed the presence of dorso Leueosiidae. Pinnotheridae and Ocypodidae (Oeypodinae). lateral knobs on the abdominal somites 2-5 as an additional This finding gives importance to Giypiograpsus in terms of feature to distinguish Plagusiinae larvae. All these larval systematica of the Grapsidae. hollowing the argumentation characters allow an easy distinction of Plagusiinae zoeae of Rice (1980). this genus should he one of the more from other grapsid zoeae, with the only exception oi advanced genera within the Grapsidae. since it shows Eitcluroiirapsiis anwricanus (see Wilson & Gore. 1980). In reduction in the number of setae (e.g. on the endopod of the the ease of the sympatric American species E. anwricanus maxilla and the basis of the first maxilliped). On the other and Plugiishi dcpressu (Fabricius. 1775). only minor hand, the possible link to the Oeypodinae and Pinnotheridae morphological features allow the idenlicication of zoeae: would rather suggest a basal position of Glyptograpsus two knob-like posterolateral protuberances of within the Grapsidae. E. anwricanus in addition to the small rounded protuberance mentioned above, the type of telson (only in the zoea I. In adult morphology, the pronounced ehelar dimorphism being of the same type in later stages), and differences in of Glxptognipsus is another character which is unique setation of the coxa! endite of the maxillae. These species among grapsid crabs (with the only exception of are even identical in characteristics which arc otherwise Ptatychirosirapsus for which no larval stages are known). A recognized as highly variable on an inlrageneric level, as for recent comparison of all American grapsid genera, based on example the number of abdominal knobs (Table I) and the mitochondrial DNA sequence, also suggests a position of number of setae or aesthelascs on the antennules. Also in these two genera clearly outside of the Varuninae and other this case, results from mitochondrial DNA sequencing grapsid subfamilies (Schubart et al.. unpublished). For a corroborate findings from larval morphology and suggest a better understanding of the systematic position of systematic position of E. anicricamts in close relationship to Glyptograpsus within the Grapsidae, the description of P. depressa and not linked to the main group of Varuninae larvae from G. jamaicen.sis (Benedict. 1892) and (Schubart e/ al., unpublished). PUttxchiroi>rapsus would be extremely useful. Table 1 and Fig. 3 give evidence for a great Results from this study strongly suggest that the grapsid morphological similarity between the first zoeae of the subfamily Varuninae as defined by Aleock (1900) needs to varunine species Euchiro^rapsus anwricanus A. Milne- be revised. Resides the possible exclusion of Edwards. 1880 and those of the subfamily Plagusiinae (see Euchirograpsits and Gi\'ptoi>rapsus. inclusion of other also Wilson. 1980). The Plagusiinae presently consist of genera presently included within the Sesarminae are 16 species belonging to two genera {Plagusia and Percnon). necessary (Schubart & Cuesta. in press) to give this Data on /oeal morphology are only available for one species subfamily a reliable phylogenetic basis. Larval morphology of Percnon and three species of Phii>n\ia (see Table 1). A alone cannot be used to determine the systematic position of prezoea of Plagusia dcpressu iubcrcidata has been the genera Euchirograpsus and Glxptograpsus. bul for the described by Rajabai (1961) (as P. d. squamosa), but is not former it clearly suggests a close similarity to the included in this tabic due to an incomplete development of Plagusiinae. while the latter. Glxptogmpsus. cannot setation patterns in that early phase. The description of presently be attributed to any other grapsid subfamily and "Plagusiinae ASM29", a lirst zoeal stage from the plankton might be of special interest for future studies of grapsid of the eastern Atlantic (Rice and Williamson. 1977). most subfamilial relationships. .1. A. CU-STA. C. IX SCIU BARI 297
Figure 3. Eitchiroi>rttpsus amcricamis. zoea I. A. lateral view: B. antenna (aTier Wilson. 1980). Pennon xibbt'si. /oea I. C. lateral view; D. antenna (alter Paula and Harlnoll. I989|. Phigitsiu il. deprcs\a. /oea I. F, lateral \ iew; F. antenna (alter Wilson and Gore. 1980). Scale bars = 0.1 mm. Figure 3. Euchirograpstis americanus, vac. I. A. vue laterals; B. aiiicnnc (d'apres Wilson. 198(1). Pennon gibbi'si. /oe 1. C. vue lute- rale ; D. antenne (d'apres Paula et Hartnoll. 1989). Plugusia Acknowledgements a la Invesiigacion)' and C.D.S. by the DFG (grant Di 479- 2/1 to Rudolf Diesel). We further want to thank Rudolf Specimens of Glxptograpsus impressus were obtained while visiting [he Smithsonian Tropical Research Institute at Diesel for kindly providing zoeae I of Hcmi^rctpsti.s Naos, Panama, where we found great support in John edwardsi from New Zealand. Jens Reimer for taking care of Christy and the local staff. Permits were granted by ovigerous females. Antonio Rodriguez for rearing larvae of 1N.RH.NA.RF Recursos Marinos. J.A.C. was financially the 1997 hatch, and Suzanne Frederieq for translating into supported by the 'Junta de Andaluct'a (Programa de Ayudas French. Peter Davie. Fernando Marques. Chang Hyun Kim 29S FIRST ZOhAI. STAGh OF GI.YPTOCRAPSUS IMPRESSES and Claude Jouin-Toulinond added valuable comments ratory. Journal of Science of the Pusan National University, when reviewing earlier versions of this manuscript. Korea. 55: 195-206. Kim C.H. & Moon D.Y. 1987. Complete larval development ol' References Hemigrapsus sinensis Rathbun (Braehyura. Grapsidae) reared in the laboratory. Korean Journal of Zoology 30: 277-291. Aikawa H. 1929. On larval forms of some Brachyiira. Records of Lebour M.V. 1944. Larval crabs from Bermuda. Zootogiea. N.Y.. Oceattographic Works of Japan. 2: 17-55. 29: 113-128. Aikawa H. 1937. Further notes on hrachyuran larvae. Records of Fee H.J. 1988. 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