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And Gecarcoidea Lalandii H. Milne

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And Gecarcoidea Lalandii H. Milne Journal of Natural History, 2002, 36, 1671–1685 First zoealstages of Epigrapsuspolitus Heller, E. notatus (Heller)and Gecarcoidealalandii H.Milne-Edwards, with remarks on zoeal morphologyof the Gecarcinidae Macleay (Crustacea: Brachyura) JOSE´ A.CUESTA†, HUNG-CHANG LIU‡and CHRISTOPH D. SCHUBART†1 †Department ofBiology, Laboratory for CrustaceanResearch, University ofLouisianaat Lafayette,Lafayette, LA 70504-2451,USA; e-mail: [email protected] ‡Department ofLife Science, National Tsing Hua University, Hsinchu, Taiwan (Accepted10 April 2001 ) Thecrab family Gecarcinidae Macleay, 1838 currently consists of 18speciesthat aregrouped in four genera. Larval data for the Gecarcinidae were only known forspecies of the genera Cardisoma Latreille,1825 and Gecarcinus Leach,1814. Inthe present paper, the rstzoeal stage of Gecarcoidealalandii H. Milne- Edwards,1837, Epigrapsuspolitus Heller,1862 and E. notatus Heller,1865 are describedand illustrated. Zoeal morphology of the Gecarcinidae is reviewed, takinginto account all previous descriptions and analysing the relationships betweenthe di Verentgenera of Gecarcinidae based on zoeal morphological characters.A seriesof typical morphological features is proposed for the zoea larvaeof this family di Verentiatingthem from the rest of the Grapsoidea. Keywords: Larvalmorphology, Gecarcinidae, Gecarcoidea , Epigrapsus, zoea. Introduction The grapsoid familyGecarcinidae has acircumtropical distribution, with many species known only from oceanic islands. The familypresently consists of 18species that aredistributed among four genera: Cardisoma Latreille,1825, Gecarcinus Leach, 1814, Gecarcoidea H.Milne Edwards,1837 and Epigrapsus Heller, 1862( Tu¨ rkay, 1970,1973, 1974; Tavares, 1991 ). At this point, wefollow Tavares( 1991)in con- sidering Discoplax A.Milne Edwards,1873 and Johngarthia Tu¨rkay,1970 subgenera of Cardisoma and Gecarcinus,respectively. Crabsincluded inthe Gecarcinidae areoften largein size and commonly referred to as‘ land crabs’, based on the terrestrial habits shown byadults of most of the 1Presentaddress: Biologie I, Universita¨tRegensburg,D-93040 Regensburg, Germany. Journalof Natural History ISSN0022-2933 print /ISSN1464-5262 online ©2002Taylor & FrancisLtd http://www.tandf.co.uk /journals DOI:10.1080 /00222930110059673 1672 J. A. Cuesta et al. species of this family.Some species canbe found severalkilometres awayfrom the coast (Gilchrist, 1988).However,no gecarcinidspecies isatrue land crab, since all of them haveto return to the seafor larvalrelease. All zoealarvae so far known aremarine planktonic, and larvaldevelopment consists of veto sixzoeal stages and one megalopalstage. Currently, larvaldata are available for only two generaand sixspecies of Gecarcinidae. Complete larvaldevelopment isknown for Cardisoma( C.)guanhumi Latreille,1825 (Costlow and Bookhout, 1968), Cardisoma( C.)carnifex (Herbst, 1794)(Kannupandi et al., 1980), Gecarcinus(G.) lateralis (Freminville,1835 ) (Willems, 1982)and Cardisoma(Discoplax) hirtipes Dana,1851 (Shokita and Shikatani, 1990). The prezoea of Cardisoma( C.)armatum Herklots, 1852has been described byCannon (1923)and the prezoea and rst zoealstage of Gecarcinus (Johngarthia)planatus Stimpson, 1860has been described byErhardt and Niaussat (1968). Previous incomplete descriptions of the rst zoealstages of Cardisoma guanhumi byMoreira (1913)and Gecarcinuslateralis byCabrera ( 1965)havenot been considered in the present study due to the availabilityof newer and more complete descriptions. This paper provides adetailed morphological description of the rst zoealstages of Gecarcoidealalandii H.Milne Edwards,1837, Epigrapsuspolitus Heller, 1862and E. notatus (Heller, 1865)based on laboratory-hatched material.For both of these genera,there wereno previous zoealdescriptions. The comparison with descriptions of other gecarcinidgenera gives new insights into possible phylogenetic relationships within this family. Materials andmethods First stagezoea larvaewere obtained from two hatches of Gecarcoidealalandii , nine hatches of Epigrapsuspolitus and two hatches of E. notatus.Ovigerous crabs of Gecarcoidealalandii werecollected in Hsiang-Chiao-Wan,Pingtung Province, Taiwan( 30May 1997 and 3August 1999by H.-C.L.). One ovigerous Epigrapsus politus wasfrom Pingtung, Taiwan( 28August 1999by H.-C.L.), another one from Hsiang-Chiao-Wan,Taiwan ( 2September 1999by H.-C.L.), and seven werecollected from MalalayangBeach, Sulawesi, Indonesia (23January 2000 by C.D.S.). Two ovigerous crabs of E. notatus werecollected from Hengchun and Hualien, Taiwan (2and 30September 1999by H.-C.L.). Ovigerous crabs weremaintained in con- tainers with natural seawater until hatching. Freshlyhatched larvaewith anactive natatory behaviour were xedin 70%ethanol. Appendages weredissected under a Wild MZ8binocular microscope, and drawingswere made using anOlympus BH-2 microscope equipped with Nomarski interference contrast and attached camera lucida. All measurements weremade byan ocular micrometer. Drawingswere based on 10larvae, and measurements on 20–30 larvae. One hatch of G. lalandii, three hatches of E. politus (one from Taiwan,and two from Indonesia) and two hatches of E. notatus weremeasured. No signicant di Verences in morphology and morpho- metry werefound between hatches of the same species. In the caseof E. politus, all drawingsare based on larvaefrom one hatch (female 4)from Sulawesi.The following measurements weremade: rostro-dorsal length (rdl )wasmeasured from the tip of the rostral spine to the tip of the dorsal spine; carapacelength (cl )from the base of the rostrum to the posterior margin;carapace width (cw) asthe distance between the tips of the lateralspines. The long natatory setae on the distal exopod segments of the rst and second maxillipedsare truncated in gures 2and 5.Descriptions Larvaldevelopment of Gecarcinidae 1673 and gures arearranged according to the standard proposed byClark et al. (1998). Maternal crabs of Gecarcoidealalandii , Epigrapsuspolitus , and E. notatus were deposited atthe National TaiwanMuseum, Taipei( TMCD3276– 3278 ). Samples of larvaeof the three species weredeposited atthe United States National Museum of Natural History, Washington, DCunder the accession number USNM 310337, 310336and 310335respectively. Results Descriptions Gecarcoidealalandii H.Milne-Edwards, 1837 Zoea I (gures 1A–C, 2A– D, 3A–C ) Dimensions. rdl: 0.54Ô 0.02mm; cl:0.33 Ô 0.01mm; cw:0.37 Ô 0.01 mm. Cephalothorax (gure 1A). Globose, smooth and without tubercles, with an anterodorsal protuberance asa carina.Dorsal spine short and slightlycurved. Rostral and lateralspines short and straight. One pair of posterodorsal setae. Anterodorsal region, posterior and ventralmargin without setae. Eyessessile. Antennule (gure 1B).Uniramous. Endopod absent. Exopod unsegmented with three aesthetascs (two long and one thin and short)and two setae. Antenna (gure 1C). Well-developed protopod almost reachingthe tip of rostral spine and bearingtwo rows of well-developed spines. Exopod elongated, reaching the middle of the protopod and bearinga group of three minute subterminal spinules, one long and one short terminal setae, and veshort terminal spines. Mandible.Endopod palp absent. Maxillule (gure 2A). Coxalendite with sixplumodenticulate setae. Basialendite with vesetae (two cuspidate and three plumodenticulate). Endopod two-segmented with one plumodenticulate seta on the proximal segment and one medial,two subterminal and two terminal plumodenticulate setae on the distal segment. Exopod absent. Maxilla (gure 2B).Coxal and basialendites bilobed with 5 1 4plumodenticulate setae. Endopod unsegmented, bilobed with one long plumodenticulate and one short simple seta on inner lobe and two long plumodenticulate setae on outer lobe. Scaphognathite with four plumose marginalsetae and one long setose posterior process. First maxilliped (gure 2C). Coxawith one seta. Basiswith 10medial setae arranged2, 2,3, 3.Endopod ve-segmented with 2,2,1, 2,5(one subterminal 1 four terminal )setae. Exopod two-segmented, with four long terminal plumose natatory setae on the distal segment. Dorsal part of basis and exopod covered with minute spinules. Secondmaxilliped (gure 2D). Coxawithout setae. Basiswith four medial setae arranged1, 1,1, 1. Endopod three-segmented with 1,1, 6 (three subterminal 1 three terminal )setae. Exopod two-segmented, with four long terminal plumose natatory setae on distal segment. Dorsal part of basis and exopod covered with minute spinules. Thirdmaxilliped . Absent. Pereiopods. Absent. 1674 J. A. Cuesta et al. Fig. 1. Gecarcoidealalandii H.Milne Edwards, 1837, Zoea I. (A) Cephalothorax,lateral view;( B)antennule;(C )antenna.Scale bars: ( A)0.1mm, ( B,C)0.05mm. Abdomen (gure 3A,B). Five abdominal somites. Somites 2–3with one pair of dorsolateral processes. Somites 3–5with conspicuous posterolateral processes. Somites 2–5 with one pair of posterodorsal setae. Pleopods absent. Telson (gure 3A–C ). Bifurcated with three pairs of stout spinulate setae on posterior margin.Along the distal part of each furcal arm two rows of minute spines. Twolateral spines on outer marginof eachfurcal arm. Larvaldevelopment of Gecarcinidae 1675 Fig. 2. Gecarcoidealalandii H.MilneEdwards, 1837, Zoea I. (A) Maxillule;( B) maxilla; (C)rstmaxilliped; ( D)second maxilliped. Scale bars: ( A,B) 0.05mm, (C, D)0.1 mm. Epigrapsuspolitus Heller, 1862 Zoea I (gures 4A–C, 5A– D, 6A–C ) Dimensions. rdl: 0.42Ô 0.02mm; cl:0.24 Ô 0.01mm; cw:0.33 Ô 0.01 mm. Cephalothorax (gure 4A). Globose, smooth and without tubercles,
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