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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 Larvae and first-stage juveniles of the American Abele, 1992 (Brachyura: ): a morphological description of two complete developments and one first zoeal stage Guillermo Guerao a; Klaus Anger b; Christoph D. Schubart c a IRTA, Unitat de Cultius Experimentals, Carretera del Poblenou Km 5.5, 43540 Sant Carles de la Ràita, Tarragona, Spain b Alfred-Wegener-Institut für Polar- und Meeresforschung, Biologische Anstalt Helgoland, Helgoland, Germany c Biologie I, Universität Regensburg, Regensburg, Germany

Online Publication Date: 01 January 2007 To cite this Article: Guerao, Guillermo, Anger, Klaus and Schubart, Christoph D. (2007) 'Larvae and first-stage juveniles of the American genus Armases Abele, 1992 (Brachyura: Sesarmidae): a morphological description of two complete developments and one first zoeal stage', Journal of Natural History, 41:29, 1811 - 1839 To link to this article: DOI: 10.1080/00222930701500431 URL: http://dx.doi.org/10.1080/00222930701500431

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Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 n n is oa stage zoeal first developments one complete and two of description morphological History Natural of Journal orsodne .Gea,IT,Uia eClisEprmnas artr e olnuK . 34 atCre ela de Carles 10.1080/00222930701500431 Sant online DOI: 43540 1464-5262 5.5 print/ISSN Km 0022-2933 Poblenou ISSN del Carretera Experimentals, Brazil, from Cultius de coast southern Unitat Ra west IRTA, to American Guerao, the G. Massachusetts Correspondence: along from and Caribbean, coast the east and Mexico American of the Gulf et the Schubart along including (see occur Sesarmidae family 2000) terrestrial al. to semi-terrestrial the of Representatives Introduction Keywords: of crab first the and larvae the sesarmid the of stages crab first and development larval Complete Abstract 1 GUERAO GUILLERMO genus American the of Armases juveniles first-stage and Larvae aoaoyclue l avlsae fu oa tgsadtemglp)adtefrtjvnl crab juvenile first the and megalopa) the and stages zoeal (four stages of stage larval All culture. and laboratory 1853) Edwards, Ra ( Germany Regensburg, HITP .SCHUBART D. CHRISTOPH ntl egln,Meesain egln,Gray and Germany, Helgoland, Meeresstation, Helgoland, Anstalt n opoercdfeecs u h eaoa fbt pce a aiyb itnuse ythe by distinguished be easily can both sternum. of the megalopae of the setation but differences, morphometric and genera the of larvae known other of those with compared are species sesarmid three Armases the of larvae the of eeomn,mglp,Ssria,zoea Sesarmidae, megalopa, development, ftecx ftefrtmxlie.Mnrdfeecswr loosre ewe h ale n the and earlier the between observed of morphologies also zoeal were The differences setation. cephalothoracic Minor of maxilliped. description first the present of coxa the of is iefrzelsae ftegenus the of stages zoeal for time first cetd2 a 2007) May 29 Accepted RA nttd utu xeietl,CreeadlPbeo m55 34 atCre ela de Carles Sant 43540 5.5, Km Poblenou del Carretera Experimentals, Cultius de Unitat IRTA, pt,Traoa pi.Eal [email protected] Email: Spain. Tarragona, `pita, pt,Traoa Spain, Tarragona, `pita, .americanum A. .roberti A. bl,19 and 1992 Abele, rae mrcnm rae iod,Amssroberti, Armases ricordi, Armases americanum, Armases bl,19 Bahua eamde:a Sesarmidae): (Brachyura: 1992 Abele, r ecie n lutae ndti o h is ie n rvosdsrpinof description previous a and time, first the for detail in illustrated and described are d asue 88 sdsrbdfrtefrttm.Temrhlgclcaatr of characters morphological The time. first the for described is 1858) Saussure, (de .ricordi A. .ricordi A. Aratus 07 12–2:1811–1839 41(29–32): 2007; , 2 .ricordi A. lrdWgnrIsiu fu Alfred-Wegener-Institut 1 .MleEwrs 87 eea hrcesaedsrbdfrthe for described are characters Several 1837. Edwards, Milne H. n diinlcaatr r nlddhr,i atclrthe particular in here, included are characters additional and , H in dad,15)fo aac eeotie from obtained were Jamaica from 1853) Edwards, Milne (H. LU ANGER KLAUS , Armases # srvsdadcmlmne.I diin h is oa stage zoeal first the addition, In complemented. and revised is 3 07Tyo Francis & Taylor 2007 seilymnt pnso h esnadtesetation the and telson the on spines minute especially , .roberti A. rPlr n ersoshn,Biologische Meeresforschung, und Polar- ¨r 2 & and 3 ilgeI Universita I, Biologie .ricordi A. rcyr,jvnl,larval juvenile, Brachyura, rae roberti Armases nyso io meristic minor show only tRegensburg, ¨t H Milne (H. Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 seAgre l 06,toeof those 2006), al. et Anger (see dl pcmn of specimens Adult methods and Material roberti eei pce ihnnabeitddvlpet n rmtecoeyrltdgenus related closely the from and development, non-abbreviated with species generic of logy for that noted stages 38) p zoeal (1992, four Abele with 2000). developments larval non-abbreviated and (A marine show species Both ricordi A. 2konseis(bl 92 im1996). Niem 1992; (Abele species known 12 hsooy reigpten,adslnt oeac ftelra tgsof stages larval the and of ecology tolerance adult salinity the and of patterns, Aspects breeding unpublished). physiology, Schubart, Herna D. (C. 1992; Trinidad from (Abele record Mexico) recent continental of exclusively (Gulf Tampico is among from but and ranges rubble species and this Yucata rocks of across to under distribution America and known among South The streams, vegetation. and freshwater Central small 1998). of among Schuh common coast and Diesel Caribbean 1992; the (Abele Surinam , Indies, from ranges West species This Bahamas, inland. metres Bermuda, hundred several Hobbs to up fringe and supralittoral the (Chace Trinidad to in species Cuba from 1992). islands, and distribution Abele known (1998) 1969; Caribbean Its Schuh characteristics). the and habitat of Diesel comprises description (see exclusively detailed areas for seepage (2006) al. and et streams Anger freshwater of banks the along he oa tgs(usae l 99,ocrigi urtdlrc ol sapeculiar a as pools rock supratidal 1995). in Diesel occurring and Schuh 1999), 1995; al. (Anger et habitat’’ (Cuesta ‘‘breeding stages zoeal three hoaohr atrs(ne ta.20) l hs pce astruhfu zoeal four through pass species these All 2006). for al. except et of stages, juvenile (Anger first the patterns and chromatophore stages larval The recently (2006). roberti most Abrunhosa A. and and (1999), Lima al. by et 1897) Cuesta (Rathbun, and (1997) Zimmermann-Ruf by 1897) Dn,15)b oacu 19)adCet n ne (2001); Anger and Cuesta and (1994) Kowalczuk by 1852) (Dana, Dı by 1897) (Rathbun, cinereum development’’. larval marine or estuarine an suggesting genus The 1817. mrcnseisaecasfe ihnfu genera: four the Presently, within coasts. tropical classified and subtropical Armases are the along species found be American can diversity high a while eprt ein [i.e. regions to temperate (Mexico) California Baja 1812 tGitnn aes(rlwy aac) h rb eetasotdaiet the to alive transported were crabs The Jamaica). (Trelawny, Waters Glistening at ´ rae roberti Armases ee edsrb o h is ietecmlt avladeryjvnl eeomn of development juvenile early and larval complete the time first the for describe we Here, h avlmrhlg of morphology larval The vrzadEad19;Dee n cu 98 cuatadDee 98 isle al. et Diesel 1998; Diesel and Schubart 1998; Schuh and Diesel 1990; Ewald and lvarez n h is oa tg of stage zoeal first the and .ricordi A. bl,19;teJmia endemic Jamaican the 1992; Abele, yHmn(94 n oto n okot(1960); Bookhout and Costlow and (1924) Hyman by aepeiul ensuid(isladShh19;Shbr n isl1998). Diesel and Schubart 1998; Schuh and (Diesel studied been previously have .Gea tal. et Guerao G. aercnl endcmne htgahcly ihabifdsrpino the of description brief a with photographically, documented been recently have Armases n eie n utml oteCrbencat fCsaRc n Panama, and Rica Costa of coasts Caribbean the to Guatemala and Belize, ´n, n opr avladjvnl etrswt hs nw rmtecon- the from known those with features juvenile and larval compare and , Armases H in dad,15)i omnsm-ersra rboccurring crab semi-terrestrial common a is 1853) Edwards, Milne (H. n a enrpre rmdycatlhbtt mn ok n ersin debris and rocks among habitats coastal dry from reported been has and .miersii A. rae roberti Armases rae ricordi Armases a n wl 16)adShbr n usa(1998); Cuesta and Schubart and (1968) Ewald and ´az eam reticulatum scoeyrltdto related closely is hc hw nabeitdmd fdvlpetwt only with development of mode abbreviated an shows which , Armases .americanum A. rae ricordi Armases Aee19) nyafwssri pce iei the in live species sesarmid few a Only 1992). (Abele ´ eecletdi 03aogtebnso h i Bueno Rio the of banks the along 2003 in collected were H in dad,15)i n ftemr terrestrial more the of one is 1853) Edwards, Milne (H. a rvosybe ecie o iespecies: five for described been previously has .americanum A. Sy 87 and 1817) (Say, Aratus eietelra n al ueiemorpho- juvenile early and larval the revise , ewe h ec n h agoezone the and beach the between rae americanum Armases ne gieae l 96,ecp o a for except 1996), al. et Aguilera ´ndez seFaiie l 05 n osssof consists and 2005) al. et Fratini (see Aratus h ‘gsaesaladnumerous, and small are ‘‘eggs the ahu,19;and 1896; Rathbun, rae cinereum Armases .MleEwrs 1837; Edwards, Milne H. .ricordi A. d asue 88 is 1858) Saussure, (de .miersii A. rae roberti Armases and Bs,1802)], (Bosc, Sesarma .angustipes A. .benedicti A. .rubripes A. (Rathbun, Aratus Say, and A. A. . Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 ctlgenmesIM 84/07adIM 44/07frtelra tgsand stages larval Cie the de for 44–49/2007 Institut of ICMD juvenile and the first 38–43/2007 ICMD of numbers Reference (catalogue of Collections (2004). Garm and (1992) Ingle followed we terminology, nytemi ifrne rmtefrtze r ecie ndti.Temrhlgcland morphological The detail. in described are zoea first the from differences main the of only stage zoeal first The Results ( Collection Zoological Lafayette al et basic Clark the (see exopod followed to descriptions endopod and Larval from proximal maxillipeds and from truncate. described the segments is are appendages of distal on armature exopod plumose to scaphognathite setal d, and distal 7c, pattern, Figure the the body In malacostracan instead. on truncated of drawn setae are and the setae illustrated plumose of fully aesthetascs five not long long between The are available. pleopods varied the were morphology as and hatches the many describe as antennules from to zoeae examined including 10, stage and width larval carapace each of carapace; the the individuals carapace. of the of the as (base margin across measured telson distance posterior was the greatest the (cl) (bt), of the to length length as margin carapace margin telson (cw) posterior juvenile, frontal basal first the the tip; and from to furcal megalopa distance margin the the to anterior For the telson the seta). of margin the outer across base of the distal line across margin the a line the posterior imaginary to from to an the exopod from exopod on antennal (fl) seta the antennal length furcal of outer the process; base protopodal the of the from of base (pl) tip margin; length spines; the carapace process rostral posterolateral protopodal from the and setae); (el) to dorsal (without spine the length rostral of the exopod of tips taken: base antennal the were the between from measurements (cl) distance length following the carapace The as setal the appendages. (rdl) stage of length the before determination larval rostro-dorsal of the and died for measurements hatch used and was but each microscope formula of field. BH-2 alive, individuals Olympus the An (10 in kept measured). individuals food were of were appropriate measurements of others and lack dissection the ethanol, to due in II, Some fixed respectively. zoea 1999, to immediately February moult 14 of successful and were days 13 following on them the hatched during females of freshwater two of with Larvae filled work. containers field small in maintained were and Montepı of beach narrow otlt.Euieadseieso ahdvlpetlsaewr rsre n70% in preserved were stage developmental and moults each for checked of were specimens larvae ethanol. the and and daily Exuviae changed was mortality. Water Lake). ( Salt three (Great isopods from sp. Larvae marine conditions. frozen same the with roberti under daily place fed took of eggs hatches were of crabs incubation and The 24 extrusion of cycle. temperature salinities dark constant at a maintained h respectively, 12 were psu, 32 they and where 0.2 (Germany), of Station Biological Marine Helgoland aetlaiasadsmlso l avlsae eedpstda h Biological the at deposited were stages larval all of samples and Parental l rwnswr aewt h i fmcocp htgah.Tenme of number The photography. microscope of aid the with made were drawings All the for used was micrometer, ocular an with equipped microscope, binocular M7 Wild A vgru eae of females Ovigerous r3 s ( psu 32 or ) .roberti A. rae roberti Armases .ricordi A. n w ace of hatches two and rae roberti Armases rae americanum Armases o(earz eio ytetidato n1 eray1999 February 11 on author third the by Mexico) (Veracruz, ´o aiiy n e al ihfehyhthdnuliof nauplii hatched freshly with daily fed and salinity, ) and rae americanum Armases .ricordi A. sdsrbdcmltl.Frtesbeun stages, subsequent the For completely. described is aveadfrtsaejvnlsof juveniles first-stage and Larvae .ricordi A. epciey n h nvriyo Louisiana of University the and respectively) , eecpue erasalsra rsigthe crossing stream small a near captured were eems-erda 24 at mass-reared were nisdlMr(SC nBarcelona in (CSIC) Mar del `ncies aaou ubrUL 4027). ULLZ number catalogue ; u ,ada riiil1 light– h 12 artificial an and C, . rae 1813 Armases u 98.Frsetal For 1998). n 5pu( psu 25 and C Idotea p) The sp.). Artemia A. Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 preypuoestearne 2 arranged 5 setae plumose sparsely with 9a). (Figure plumose bilobed maxilliped four with First endite (exopod) process. posterior Basial Scaphognathite setose respectively. long 7a). Figure lobe, and outer setae in and marginal inner 2 (detail with on bilobed, lobe setae one unsegmented, distal Endopod and setae. on plumodenticulate segment, tooth proximal marginal on segment. 7a). (Figure distal seta Maxilla the with one on and setae plumodenticulate) with plumodenticulate two terminal two-segmented, and four and cuspidate Endopod subterminal (three setae teeth. five two with endite Basial setae. 5a). (Figure Maxillule Mandible. ipeste(n og n eimszd n w iue.pl/el terminal minute). four two with and elongated, medium-sized, Exopod one sizes. long, different (one of setae (8–11) simple spines of number unequal 4a). (Figure Antenna seta. terminal one and aesthetascs terminal sessile. 3a). Eyes (Figure setae. Antennule without posterodorsal margin on ventral setae of and Pair straight, Posterior absent. present, regions. spines spine Lateral anterodorsal protopod. Rostral and antennal posteriorly. to curved length in strongly similar developed, well present, spine 1a). (Figure Carapace lms ea ragd1 arranged setae 10a). (Figure plumose maxilliped subterminal Second long (one four 5 with 2, segment 1, distal setae. 2, two-segmented; natatory 2, plumose Exopod with setae. five-segmented, plumodenticulate Endopod terminal) side. proximal outer Size. I Zoea Description I. Table hr maxilliped. setae. Third natatory plumose long four with segment distal two-segmented; of individuals with differences meristic 1814 rdl 5 .Gea tal. et Guerao G. 0.85 apasn;mlradicsrpoesswl developed. well processes incisor and molar absent; Palp ¡ .2m;cl mm; 0.02 Absent. oa nieblbd ih5 with bilobed, endite Coxal iia nsz orsrlsie rtpdlpoeswt w osof rows two with process Protopodal spine. rostral to size in Similar lbs,sot,wt iueatrltrlpoueac.Dorsal protuberance. anterolateral minute with smooth, Globose, xpdadeio ea bet oa niewt i plumose six with endite Coxal absent. setae epipod and Exopod nrmu.Edpdasn.Eoo nemne ihfour with unsegmented Exopod absent. Endopod Uniramous. + 1 + oawt n preypuoest.Bsswt 0medial 10 with Basis seta. plumose sparsely one with Coxa 5 1 + 0.52 .Edpdtresgetd ih0 ,6ste Exopod setae. 6 1, 0, with three-segmented, Endopod 1. oawtotste ai ihfu eilsparsely medial four with Basis setae. without Coxa + 2 ¡ rae roberti Armases + 3 Fgrs1–16) (Figures .3mm. 0.03 + ninrsd,adamto ogmcorcieon microtrichiae long of mat a and side, inner on 3 .ricordi A. + and lmdniuaesteadwt a with and setae plumodenticulate 3 .americanum A. + ogplumodenticulate long 3 5 2.0–2.2. r rsne in presented are + four + 4 oa2 Zoea oa4 Zoea 1 Maxilliped al .Mrhlgcldfeecsaoglra tgso h genera the of stages larval among differences Morphological I. Table oa3 Zoea development). oa1 Zoea aaaev s d22n dn 1 nd nd nd 2 2 nd 2 (s) vm Carapace aaaed s d22n d00 8 0 6 nd nd nd nd 2 10 2 10 nd nd 2 8 (s) ds Carapace (s) ad Carapace l(m 0.34 (mm) cl l(m 0.47 (mm) cl uc ms)2111n dn nd nd 6 nd nd 5 2 nd nd 5 nd nd nd nd 5 1 1 nd 6 1 1 2 6 1 1 6 2 0.70 2 1 2 5 (mm) rdl (mlsp) Furca Telson 4 (s) Coxa 1 Maxilliped (s) Coxa Maxillule (a Antennule (s) ad Carapace aaaed s d22n d00 6 4 0 6 nd nd nd 5 5 2, 2, 3, 2, 5 2, 1, 2, 2, nd nd 5 5 2, 2, 2, 2, 5 2 2, 2, 2, 2, 8 5 5 2, 2, 3, 2, 2 5 2, 8 2, 3, 2, 5 nd nd 0.92 nd nd 5 2, 2, 3, 2, 0 8 (mm) rdl 5 (s) Endopod (s) ds Carapace (s) ad Carapace (s) vm Carapace l(m 0.41 (mm) cl boe o s d11n d00 6 0 nd nd nd nd nd 1 4 1 4 nd nd 0.79 1 4 (mm) rdl (s) so1 Abdomen (s) ad Carapace l(m 0.30 (mm) cl d m)0.57 (mm) rdl + )44555455 5 4 5 5 5 4 4 s)

Downloaded By: [Guerao, Guillermo] 2006a) al. et At: 06:40 22 October 2007 (Cuesta ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ .2n 0.72 nd 0.02 .2n 0.97 nd 0.02 .3n 1.12 nd 0.03 .4n 1.50 nd 0.04 .3n 0.85 nd 0.03 .3n 1.45 nd 0.03 .20.45 0.02 .40.73 0.04 americanum (present Armases paper) ¡ ¡ .20.52 0.02 .30.85 0.03 (present Armases paper) roberti ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ .20.58 0.02 .50.91 0.05 .40.95 0.04 .71.45 0.07 .30.73 0.03 .30.49 0.03 .41.15 0.04 .20.70 0.02 Armases and (present Armases paper) ricordi Aratus ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ .2n dn 0.53 nd nd nd 0.02 .3n dn 0.71 nd nd nd 0.03 .2n dn 0.97 nd nd nd 0.02 .4n dn 1.30 nd nd nd 0.04 .2n dn 0.66 nd nd nd 0.02 .20.48 0.02 .3n dn 1.21 nd nd nd 0.03 .30.72 0.03 o considering not , n usa1998) Cuesta and Da n Ewald and (Diaz 98 Schubart 1968; rae ricordi Armases ¡ ¡ .10.37 0.01 .20.63 0.02 .benedicti A. n usa1998) Cuesta and Da n Ewald and (Diaz rae rubripes Armases 98 Schubart 1968; iacrt ecito)and description) (inaccurate ¡ ¡ .2n 0.41 nd 0.02 .3n 0.78 nd 0.03 cinereum n Bookhout and Armases 1960) (Costlow .miersii A. ne 2001) Anger Cet and (Cuesta angustipes (abbreviated Armases ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡

0.02 0.02 0.03 0.04 0.02 0.03 0.03 0.04

aveadfrtsaejvnlsof juveniles first-stage and Larvae rae 1815 Armases al .( I. Table Maxilla Maxillule Antenna Antennule Megalopa 1 Maxilliped ailpd2 Maxilliped 1 Maxilliped Maxilla xpd()1 d1 ,51 ,50 ,6 1, 5 0, 5 1, 8 4, 1, 0, 6 6, 5 1, 1, 0, 6 3, 1, 0, 5 1, 6 3, 1, 0, 5 1, 7–8 4–5, 1, 1, 6–7 3, 1, 0, nd nd 5 1, 7 3, 1, 0, (s) Exopod (s) Endopod ailedt s 7n 61 81 71 15 11 12 3 6, 9 0, 5 6, 0, 17 11 nd 18 11 3 1, 5, 1, 2, 0, nd 2 1, 2, 18 1, 2, 0, 11–12 3 1, 4, 1, 2, 0, 3 7, 16–18 0, 11–12 3 1, 5, 1, 2, 0, 3 1, 3 nd 5, 6, 1, 0, nd 2, 0, 3 1, 5, 1, 2–3, 0, 8 nd 17 nd 9 (s) endite 3 Coxal 1, 5, 3 1, 6, 2, 0, 0, (s) endite Basial (s) endite Coxal (s) Flagellum (a) Exopod w(m 0.46 (mm) cw xpd()1 d9999910 nd 9 nd 9 20–21 nd 23 6 2, 9 2, 3, 2, nd 6 2, 1, 3, 2, 20 6 9 2, 2, 2, 2 2, 6 2, nd 2, 3, 2, 9 2 6 2, 2, 3, 2, 22–24 6 2, nd 2, 3, 2, nd 18–21 0.69 nd 10 1 nd 6 2, 2, 3, 2, (mm) cl (s) Exopod 23 (s) Endopod (s) Coxa (s) Scaphognathite ppd()4n 5 5 4 3, 7 5 3, 39 30 4 5 3, 35 4 3, 6 7 12 8–9 3–4, 45 7 11 6 3–4 3, 36–39 nd nd nd nd 36–39 4 3 10 3, 9 nd Coxa (s) Epipod (s) Exopod 33 (s) Basis (s) Coxa (s) Scaphognathite ailedt s 8 (s) endite Basial ailedt s 7 (s) endite Basial oa nie()6 (s) endite Coxal + ai s d040002 0 0 0 4 0 nd 0 (s) basis Continued. )

Downloaded By: [Guerao, Guillermo] 2006a) al. At:et 06:40 22 October 2007 (Cuesta Aratus pisonii ¡ ¡ + + + + 4nd12 5nd7 5nd7 4nd7 .3n 0.85 nd 0.03 .5n 1.18 nd 0.05 americanum (present Armases paper) (present Armases paper) roberti + + ¡ ¡ + – 6 5–6 – 7 4–5 + 7–9 7 .70.75 0.07 .81.10 0.08 10–11 5 (present Armases paper) ricordi ¡ ¡ + + + + + 31 11 8 9 11 5813 6 88 66 – 6 7–8 511 7 .4n dn 0.48 nd nd nd 0.04 .6n dn 0.66 nd nd nd 0.06 + 511 n usa1998) Cuesta and Da n Ewald and (Diaz 98 Schubart 1968; rae ricordi Armases + + + 66 66 411 n usa1998) Cuesta and Da n Ewald and (Diaz rae rubripes Armases 98 Schubart 1968; + + + + 66 67 47 38 cinereum n Bookhout and Armases 1960) + + + + (Costlow 66 78 511 56 ne 2001) Anger Cet and (Cuesta angustipes Armases ¡ ¡

+ + + +

5 7 4 0.03 0.05 5

1816 .Gea tal. et Guerao G. ,ashtss d neoosl s oslsie lp iueltrlsie d odt;p,poess ,ste o oie m eta margin. ventral vm, somite; so, setae; s, processes; pr, data; no nd, spine; lateral minute mlsp, spine; dorsal ds, anterodorsal; ad, aesthetascs; a, 4 3, Pereiopods 3 Maxilliped al .( I. Table rpd()1 d1 ,71 ,61 ,7 1, nd 7 1, nd nd 6 1, nd nd 7 1, nd nd 7 1, nd 6 4, 1 3, 8, 10, 7 1, 5 2, 3, 6, 11–12 7, 6–7 3–4, 8–9, 8, 1 nd 6 3, 8 4, 9, 8, 13–15, 14–15, nd 6 7 1, 4, 4, nd 8, 8, nd 7 4, 3–4, 8, 8–9, 11 12, 12, 14, nd nd 14, 14, (s) 12, Uropod 11, 6 (s) Pleopods 6 4, 3, 8, 8, (s) Abdomen (pr) Coxa (s) Sternum (s) Endopod Continued. )

ta.2006a) Downloaded al. et By: [Guerao, Guillermo] At: 06:40 22 October 2007 (Cuesta Aratus pisonii 4 4 14, americanum (present Armases paper) d7 2 4 6 14, 16, 14, 12, 7, nd 31,11–12 13–14, (present Armases paper) roberti 4 ,1,1,1,14, 14, 14, 12, 7, 5 4 3 01,1,1,1 3 3 3 11,1,1,1 3 6 7 15 17, 16, 13, 11 13, 13, 13, 11 13, 13, 13, 11 13, 13, 13, 10 13, 14, 15, (present Armases paper) ricordi 4 n usa1998) Cuesta and Da n Ewald and (Diaz 98 Schubart 1968; rae ricordi Armases dn d7 0 0 10, 10, 10, 7, nd nd nd n usa1998) Cuesta and Da n Ewald and (Diaz rae rubripes Armases 98 Schubart 1968; cinereum n Bookhout and Armases 1960) (Costlow ne 2001) Anger Cet and (Cuesta angustipes

Armases

2 4 12,

aveadfrtsaejvnlsof juveniles first-stage and Larvae rae 1817 Armases Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 1818 iue1. Figure rae roberti Armases .Gea tal. et Guerao G. aaae aea iw a oaI b oaI;()ze I;()ze V cl a:04mm. 0.4 bar: Scale IV. zoea (d) III; zoea (c) II; zoea (b) I; Zoea (a) view. lateral carapace, , Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 rcse.Smts35wt otrltrlpcse.Smts25wt arof e). pair 15a, e, a 14a, (Figures Telson with 2–5 Somites pocesses. Pleopods. posterolateral with setae. simple 3–5 posterodorsal Somites processes. 15a). 14a, (Figures Abdomen mm. 0.5 bars: Pereiopods. Scale margin. anterolateral the of detail with view, 2. Figure Size. two and margin II fl/bt Zoea outer arms. on furcal spine of scale-like margins dorsal minute on A spines margin. small posterior arrow-shaped on setae serrulate rdl rae roberti Armases 5 Absent. 1.12 Absent. ¡ .4m;cl mm; 0.04 aaae a eaoa aea iw b eaoa oslve;()frtca,dorsal crab, first (c) view; dorsal megalopa, (b) view; lateral Megalopa, (a) carapace. , 5 iesmts oie n ihapi fdorsolateral of pair a with 3 and 2 Somites somites. Five 0.72 iuct,fralre lgtydvretwt he ar of pairs three with divergent slightly large, furca Bifurcate, ¡ .2mm. 0.02 aveadfrtsaejvnlsof juveniles first-stage and Larvae 5 1.5–1.7. rae 1819 Armases Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 cl a:00 mm. 0.05 bar: Scale unchanged. maxilliped. First unchanged. 7b). (Figure Maxilla unchanged. Otherwise present. 5b). (Figure Maxillule Mandible. 3. Figure 1820 nen Fgr 4b). (Figure Antenna 3b). (Figure Antennule plumose sparsely (one setae unchanged. two with 1b). (Figure margin Carapace rae roberti Armases .Gea tal. et Guerao G. Unchanged. xpddsa emn ihsxln lms aaoyste Otherwise setae. natatory plumose long six with segment distal Exopod nenl.()Ze ;()ze I c oaII d oaI;()mglp;()frtcrab. first (f) megalopa; (e) IV; zoea (d) III; zoea (c) II; zoea (b) I; Zoea (a) antennule. , cpontiewt ih (3 eight with Scaphognathite noo u rsn.pl/el present. bud Endopod Unchanged. ailedt ihsvnpuoeste xpdpuoeseta plumose Exopod setae. plumose seven with endite Basial w ar fatrdra ea.Ec lateroventral Each setae. anterodorsal of pairs Two + 5 n aps) yssakd Otherwise stalked. Eyes pappose). one + .–..Ohrieunchanged. Otherwise 2.2–2.4. )puoemria ea.Otherwise setae. marginal plumose 5) Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 esn(iue1b 15b). 14b, (Figure Telson 15b). 14b, (Figures Abdomen Pereiopods. maxilliped. Third unchanged. maxilliped. Second mm. 0.05 bars: Scale megalopa. palp, Mandibular (g) crab. 4. Figure rae roberti Armases rsn sudfeetae buds. undifferentiated as Present rsn sudfeetae buds. undifferentiated as Present xpddsa emn ihsxln lms aaoyste Otherwise setae. natatory plumose long six with segment distal Exopod af nen:()ze ;()ze I c oaII d oaI;()mglp;()first (f) megalopa; (e) IV; zoea (d) III; zoea (c) II; zoea (b) I; zoea (a) Antenna: (a–f) . fl/bt is oiewt n i-oslst.Ohrieunchanged. Otherwise seta. mid-dorsal one with somite First 5 .–..Ohrieunchanged. Otherwise 1.6–1.7. aveadfrtsaejvnlsof juveniles first-stage and Larvae rae 1821 Armases Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 tews unchanged. Otherwise 3c). (Figure setose Antennule sparsely (two setae five with margin unchanged. ventral Otherwise Each spine. dorsal the 1c). (Figure Carapace 1822 Size. III Zoea iue5. Figure rdl 5 .Gea tal. et Guerao G. 1.45 rae roberti Armases ¡ .4m;cl mm; 0.04 orpiso neooslste n aro ea ert h aeof base the to near setae of pair One setae. anterodorsal of pairs Four xpdwt ortria eteac n n iuetria seta. terminal minute one and aesthetascs terminal four with Exopod aill.()Ze ;()ze I c oaII d oaI.Saebr .5mm. 0.05 bar: Scale IV. zoea (d) III; zoea (c) II; zoea (b) I; Zoea (a) maxillule. , 5 0.85 ¡ .3mm. 0.03 + he pappose). three Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 ea) pl/el setae). 4c). (Figure Antenna tews unchanged. Otherwise maxilliped. unchanged. Otherwise Second setae. natatory plumose long eight with segment distal 9b). (Figure maxilliped First unchanged. Otherwise setae. marginal 7c). (Figure Maxilla unchanged. 5c). (Figure Maxillule Mandible. iue6. Figure Unchanged. 5 .–..Ohrieunchanged. Otherwise 2.6–2.9. rae roberti Armases xpddsa emn ihegtln lms aaoysetae. natatory plumose long eight with segment distal Exopod ailedt ih5 with endite Basial noo u lnae,a oga xpdo lgtylne (without longer slightly or exopod as long as elongated, bud Endopod ppdst rsn.Bsa niewt ih ea.Otherwise setae. eight with endite Basial present. seta Epipod noo emns2ad3wt n diinlst.Exopod seta. additional one with 3 and 2 segments Endopod aill.()Mglp;()frtca.Saebr .5mm. 0.05 bar: Scale crab. first (b) Megalopa; (a) maxillule. , aveadfrtsaejvnlsof juveniles first-stage and Larvae + ea.Sahgaht ih1–3plumose 12–13 with Scaphognathite setae. 5 rae 1823 Armases Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 1824 I,wt ealo h itllb ftecxledt;()ze V cl a:00 mm. 0.05 bar: Scale IV. zoea (d) endite; coxal the of lobe distal the of detail a with III, 7. Figure rae roberti Armases .Gea tal. et Guerao G. ail.()Ze ,wt ealo h itllb ftecxledt;()ze I c zoea (c) II; zoea (b) endite; coxal the of lobe distal the of detail a with I, Zoea (a) maxilla. , Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 iue8. Figure rae roberti Armases ail.()Mglp;()frtca.Saebr .5mm. 0.05 bar: Scale crab. first (b) Megalopa; (a) maxilla. , aveadfrtsaejvnlsof juveniles first-stage and Larvae rae 1825 Armases Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 1826 rb cl as . mm. 0.1 bars: Scale crab. 9. Figure rae roberti Armases .Gea tal. et Guerao G. is ailpd a oaI b oaII noo;()ze V d eaoa e first (e) megalopa; (d) IV; zoea (c) endopod; III, zoea (b) I; Zoea (a) maxilliped. first , Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 . mm. 0.1 eeoos(iue12a). (Figure Pereiopods maxilliped. Third 10. Figure oie –,edpd bet tews unchanged. Otherwise absent. endopods 2–5, somites 15c). 14c, (Figures Abdomen rae roberti Armases iaos unsegmented. Biramous, eodmxlie.()Ze ;()ze V c eaoa d is rb cl bars: Scale crab. first (d) megalopa; (c) IV; zoea (b) I; Zoea (a) maxilliped. second , eeoosusgetd hlpd bilobed. Chelipeds unsegmented. Pereiopods oiesxpeet ihu ea.Pepdbd rsn on present buds Pleopod setae. without present, six Somite aveadfrtsaejvnlsof juveniles first-stage and Larvae rae 1827 Armases Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 1828 Size. IV Zoea 15c). 14c, (Figures Telson rdl iue11. Figure 5 .Gea tal. et Guerao G. 1.50 ¡ rae roberti Armases .7m;cl mm; 0.07 fl/bt hr ailpd a eaoa b is rb cl a:01mm. 0.1 bar: Scale crab. first (b) Megalopa; (a) maxilliped. third , 5 5 0.97 .–..Ohrieunchanged. Otherwise 2.4–2.6. ¡ .5mm. 0.05 Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 nenl Fgr 3d). (Figure Antennule plumose sparsely five to (four eight 1d). (Figure Carapace esn(iue 4,15d). 14d, (Figures Telson unchanged. Otherwise present. buds endopod elongated, 15d). 14d, (Figures Abdomen 12b). (Figure Pereiopods maxilliped. Third unchanged. Otherwise 10b). (Figure maxilliped Second Otherwise setae. natatory plumose long nine with unchanged. segment distal Exopod seta. subterminal 9c). (Figure maxilliped First unchanged. Otherwise setae. plumose marginal 18–21 with Scaphognathite 7d). (Figure Maxilla unchanged. 5d). (Figure Maxillule pl/el setae. minute without Exopod process. spinous 4d). (Figure Antenna aesthetascs. iue12. Figure ppdnwpeet rudimentary. present, now Epipod rae roberti Armases oa niewt 7 with endite Coxal noo w-emne,lne,rahn ideo h eghof length the of middle reaching longer, two-segmented, Endopod oa niewt ee ea.Bsa niewt 1ste Otherwise setae. 11 with endite Basial setae. seven with endite Coxal iepiso neooslste ahvnrlmri ihsvnto seven with margin ventral Each setae. anterodorsal of pairs Five xpdwt n aa ea n he utria n orterminal four and subterminal three and seta, basal one with Exopod hlpd n eeoos25sihl segmented. slightly 2–5 pereiopods and Chelipeds fl/bt oawt w ea.Ffhsgeto noo ihadditional with endopod of segment Fifth setae. two with Coxa is oiewt he ogmddra ea.Pepdbuds Pleopod setae. mid-dorsal long three with somite First eeoos15 a oaII b oaI.Saebr:01mm. 0.1 bars: Scale IV. zoea (b) III; Zoea (a) 1–5. pereiopods , 5 xpddsa emn ih1 ogpuoenttr setae. natatory plumose long 10 with segment distal Exopod .–..Ohrieunchanged. Otherwise 2.2–2.4. + he aps)ste tews unchanged. Otherwise setae. pappose) three aveadfrtsaejvnlsof juveniles first-stage and Larvae + – ea.Bsa niewt 7 with endite Basial setae. 4–5 5 2.7–2.9. rae 1829 Armases + – setae. 5–6 Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 1830 Size. Megalopa (j) 3; pereiopod c–i) (a, 13. Figure bet xpdtresgetd ih0 ,3ashtssad0 ,2ste respectively. setae, 2 2, 0, and aesthetascs 3 6, 0, with three-segmented, Exopod absent. 3d). undulations. (Figure Antennule prominent lateral and carina medial longitudinal with and b). 2a, (Figure Carapace cl 5 1.18 .Gea tal. et Guerao G. .roberti A. ¡ rae roberti Armases .8m;cw mm; 0.08 is rb atl eeoo .Saebr:03m ac;005m dg ) .2m (i). mm 0.02 j); (d–g, mm 0.075 (a–c); mm 0.3 bars: Scale 5. pereiopod dactyl, crab, first , eucetresgetd ih3 ,1ste epciey Endopod respectively. setae, 1 1, 3, with three-segmented, Peduncle ogrta ra.Rsrmvnrlydfetdwt eincleft, median with deflected ventrally Rostrum broad. than Longer eaoa a tru;()pripd1 dh atl eeoos25 i coxa, (i) 2–5; pereiopods dactyl, (d–h) 1; pereiopod (c) sternum; (a) megalopa: , 5 0.85 ¡ .7mm. 0.07 Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 emn ihtredsa lms ea,dsa emn ihtret orln terminal long four to three proximal with two-segmented, segment Exopod distal setae. setae, five plumose setae. with plumose distal Endopod three setae. with 11 segment with endite setae. Basial 9d). (Figure lateral with posterior maxilliped one Scaphognathite First and none. anterior or two and seta, setae subterminal marginal one plumose with 36–39 unsegmented, Endopod setae. 8a). (Figure Maxilla 18(16 aill Fgr 6a). (Figure Maxillule respectively. 4g). (Figure setae, Mandible 3 1, 5, 1, 2–3, 0, with six-segmented, 4d). (Figure Antenna I zoea of setae serrulate and furca 14. Figure + )ste noo nemne,wt w rxmladfu itlsetae. distal four and proximal two with unsegmented, Endopod setae. 2) rae roberti Armases boe,dra iw a oaI b oaI;()ze I;()ze V e ealo the of detail (e) IV; zoea (d) III; zoea (c) II; zoea (b) I; Zoea (a) view. dorsal abdomen, , oa nieblbd ih12 with bilobed, endite Coxal eucetresgetd ih0 ,1ste epciey Flagellum respectively. setae, 1 1, 0, with three-segmented, Peduncle aptosgetd ihfu ea ndsa segment. distal on setae four with two-segmented, Palp oa niewt 11 ea.Bsa niewt 16(15 with endite Basial setae. 11–12 with endite Coxal . cl as . mm. 0.1 bars: Scale ppdwt ee ogste oa niewt i setae. six with endite Coxal setae. long seven with Epipod aveadfrtsaejvnlsof juveniles first-stage and Larvae + ea.Bsa nieblbdwt 7 with bilobed endite Basial setae. 5 rae 1831 Armases + 1)– + 7 Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 rcs ah atlso it eeoo ihtreln emnlsteadoeshort medially. one interrupted and sutures setae 13a). terminal (Figure Sternum long three with pereiopod fifth spine. of terminal Dactylus each. process 15. Figure 1832 ifrnitd ihnn ea.Edpdfv-emne,wt –,8 –,4 setae, 7 13c–i). (Figure 4, Pereiopods segment setae. distal 3–4, natatory and 8, plumose seta terminal 8–9, one long with with four segment five-segmented, with proximal Endopod two-segmented, Exopod setae. respectively. nine with plumose differentiated, 11a). (Figure terminal maxilliped five Third with segment two- distal Exopod seta, respectively. setae, one 6–7 with setae. 3, segment 1, 0, proximal with segmented, four-segmented Endopod seta. without 10c). (Figure maxilliped Second mm. 0.1 bars: Scale I. zoea of furca rae roberti Armases .Gea tal. et Guerao G. boe,ltrlve.()Ze ;()ze I c oaII d oaI;()dti fthe of detail (e) IV; zoea (d) III; zoea (c) II; zoea (b) I; Zoea (a) view. lateral abdomen, , ailpd n hlpdsentsfsdwt ih ea.Alsternal All setae. eight with fused sternites cheliped and Maxillipeds l rilswl ifrnitd eeoos3ad4wt n coxal one with 4 and 3 Pereiopods differentiated. well articles All ppdeogtdwt 5ln ea.Cx n ai not basis and Coxa setae. long 15 with elongated Epipod ppdrdmnay oaadbssntdifferentiated, not basis and Coxa rudimentary. Epipod Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 loos(iue16e–h). (Figure pleopods. biramous Pleopods of pair a with each 2–5 Somites respectively. b). 16a, (Figure Abdomen mm. 0.1 bars: Scale uropod. (i) 1–4; pleopods (e–h) characters; zoeal with 16. Figure emn ihsvnln lms ea,respectively. setae, plumose long seven with segment setae, 16i). natatory (Figure plumose Uropods marginal long 11–12 13–14, 13–15, respectively. 14–15, with unsegmented rae roberti Armases eaoa a boe,dra iw b boe,ltrlve;()tlo;()telson (d) telson; (c) view; lateral abdomen, (b) view; dorsal Abdomen, (a) megalopa. , w-emne nsmt ,poia emn ihoe n distal and one, with segment proximal 6, somite on Two-segmented noo nemne,wt w emnlhos exopod hooks, terminal two with unsegmented, Endopod i oie rsn.Smts16wt ,1,1,1,1,4setae, 4 14, 16, 14, 12, 7, with 1–6 Somites present. somites Six aveadfrtsaejvnlsof juveniles first-stage and Larvae rae 1833 Armases Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 1834 iia n ge elwt e fegtcaatr eie o eamdlra yCuesta by larvae sesarmid for defined characters eight of set a with well agree and similar of morphologies zoeal The Discussion Abdomen. setae. terminal long without pereiopod 13c–j). (Figure Pereiopods setae, setae. 5 segment plumose 5, distal terminal 6–7, and long setae 16, eight five 20, with with segment with proximal five-segmented two-segmented, Exopod Endopod respectively. setae. 18–21 with undifferentiated orsgetdwt ,1 ,9ste epciey xpdtosgetd proximal two-segmented, 11b). setae. (Figure plumose Exopod terminal maxilliped respectively. five Third with setae, segment distal 9 and 5, setae eight 1, with segment 2, setae. proximal with plumose two-segmented, terminal four-segmented 10d). long Exopod (Figure four maxilliped with setae. Second segment five distal setae, with two to Endopod zero setae. with segment 20–21 with endite 9e). (Figure maxilliped marginal First plumose 48 with setae. lateral Scaphognathite eight setae. and without setae unsegmented, Endopod setae. ail Fgr 8b). (Figure Maxilla setae. distal three and proximal two with unsegmented, 6b). (Figure Maxillule respectively. setae, Mandible. 3 4, 1, 2, 0, with five-segmented, setae. flagellum 2–3 0, 4f). (Figure and Antenna respectively, aesthetascs, 3–4 0, with two-segmented, Exopod absent. third and largest 3f). (Figure first Antennule teeth, three with margins Anterolateral width. smallest. carapace of half one 2c). (Figure Carapace Size. juvenile First setae. ventral two with and setae six with surface d). c, 16a, (Figure Telson cl 5 1.35 .Gea tal. et Guerao G. i oie rsn ihpepd eue,wtotsetae. without reduced, pleopods with present somites Six aptosgetd ihfv ea ndsa segment. distal on setae five with two-segmented, Palp ¡ .2m;cw mm; 0.02 oa nieblbdwt 13 with bilobed endite Coxal eucetresgetd ih2 ,1ste epciey Usually, respectively. setae, 1 1, 2, with three-segmented, Peduncle oa niewt 4ste ailedt ih1–0ste Endopod setae. 19–20 with endite Basial setae. 14 with endite Coxal ogrta ra,qarnua.Fotlrgo ra measuring broad region Frontal quadrangular. broad, than Longer eucetresgetd ih6 ,1ste epciey Endopod respectively. setae, 1 1, 6, with three-segmented, Peduncle otro agnrudd ihoeo w lms ea.Dorsal setae. plumose two or one with rounded, margin Posterior eeoos3ad4wtotcxlpoess atlso fifth of Dactylus processes. coxal without 4 and 3 Pereiopods .roberti A. ppdwt 1ln ea.Cxledt ih1 ea.Basial setae. 12 with endite Coxal setae. long 11 with Epipod 5 1.17 ppdeogtdwt 0ln ea.Cx n basis and Coxa setae. long 20 with elongated Epipod oaadbssudfeetae,wtotste Endopod setae. without undifferentiated, basis and Coxa ¡ , .ricordi A. .2mm. 0.02 n h is tg of stage first the and + ea.Bsa nieblbdwt 9 with bilobed endite Basial setae. 5 .americanum A. r very are + 8 Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 n ea fdfeetszs h xpdbighl h egho h rtpdadthe and protopod the of length 2 rows; with the two endopod in half distributed maxillar spines number) being bilobed small in terminal a exopod unequal with (4) (usually the exopod spines sizes, well-developed antennal I with an zoea different protopod a (3) (2) of setae; spines; setae anterodorsal lateral without of and carapace pair a a (1) show: with species stage described All (2006b). al. et neiradmr eta hntehbta oiino h aea pnsi other in spines lateral the of position habitual the than ventral more and anterior of stages zoeal 2006a). the for al. described et (Cuesta maxilliped first the of coxa the ecitoso eamd u otevr ml ieo h pns pn sas rsn on present also larval is spine many A in spines. of omitted the stages of zoeal been size all has small of furca very feature telson the the This to due 2006b). sesarmids 2006a, of al. descriptions et (Cuesta Sesarmidae III), n eai oa tgsIIIadtostei oaI.Frcmaio,aprevious a comparison, For IV. zoea in setae two and genus I–III related stages closely the zoeal of description in seta one h ifrne r:acrpc ihltrlcrpc pns nedpdo h ail with maxilla the of endopod an spines, genera carapace the lateral of with 2 species carapace other a all are: to differences only The not differences, striking of Armases number a of of to al. due pattern descriptions doubtful, the et many to Cuesta in identical omitted 2004; be been to al. of has found carapace et larger the was seta of (Guerao it behind this setation hidden The Sesarmidae that zoeae. be possible family first sesarmid can is the it it within of because Therefore, common one in 2006b). visible, species, similar be easily this is to situation In not The five. seems and 5a). of Figure small instead (see very setae, setae six proximal is bears setae maxillule six the of the endite coxal the that (Dı descriptions previous for described is maxilliped genus first the the of of stages coxa zoeal the all of for setation time The first the I. Table in shown are species 1 zoeal with last basis maxilliped the second that a (and (6) 5 so 6; and development, 2, 2 2, through segments 3, while each 2, zoea, of seta first setation the endopod one in an acquire 5 have 2, stages 3) 1, segment 2, 2, also of often setation endopod an and setae nyoest;smt fze I n Vwt n ea iaadArnoa(2006) Abrunhosa and Lima seta. one with and IV The with incomplete and II abdomen. zoea III is of and zoea 5 of spines figures, and of lateral carapace 4 6 the somites the that the somite setae; suggest without from seta; of I one zoea concluded setation of only 5 setation, omitted the somite been abdominal abdominal example have contradictory: characters on for other Many description, information setae. three the with maxilliped from second the of basis rtbrne ecie for described protuberances h oiino h rtbrne,as rsn in present also protuberances, the of position the arms furcal size. and varying of with development telson spinules zoeal (8) of stage; throughout on rows last margin the found dorsal in five only posterior two 1 comprising processes somite with on stage on dorsolateral setae setae zoeal with middorsal first serrulate somites, three the with three six of and stage 3, pleon and zoeal the 2 last (7) somites the 6; of 1, 0, that of somites, setation endopod and setae + iuesaelk pnso h ue agno h esnfrahv eetybeen recently have furca telson the of margin outer the on spines scale-like Minute h eetlra ecito of description larval recent The of stages zoeal the of description current The h eitcadmrhmti ifrne ftedsrbdzelsae of stages zoeal described the of differences morphometric and meristic The ea,a noo ftefrtmxlie ftefrtze ih2 ,2 ,5ste n a and setae, 5 1, 2, 2, 2, with zoea first the of maxilliped first the of endopod an setae, 2 .cinereum A. and Aratus ZI), (Z u lot h oa rudpteno h eamde(e above). (see Sesarmidae the of pattern ground zoeal the to also but , a n wl 98 cuatadCet 98.W observed We 1998). Cuesta and Schubart 1968; Ewald and ´az .rubripes A. .rubripes A. .benedicti A. rtspisonii Aratus rae benedicti Armases .roberti A. ZI,admn te pce eogn othe to belonging species other many and I), (Z + and Aratus aveadfrtsaejvnlsof juveniles first-stage and Larvae ea;()afrtmxlie ai ih2 with basis maxilliped first a (5) setae; 3 oa r oooost h ml anterolateral small the to homologous are zoeae .ricordi A. Armases , , .roberti A. .ricordi A. rae angustipes Armases eotdfralfu oa tgsoest on seta one stages zoeal four all for reported .ricordi A. yLm n buhs 20)appears (2006) Abrunhosa and Lima by .roberti A. .ricordi A. TbeI): (Table yDı by n h is oaof zoea first the and seFgr ;TbeI). Table 1; Figure (see .roberti A. a n wl 16) However, (1968). Ewald and ´az ifr nsm etrsfrom features some in differs a o rvosydescribed; previously not was and .roberti A. , .miersii A. .americanum A. and .americanum A. and rae 1835 Armases , .americanum A. .ricordi A. .ricordi A. smore is , Armases + + 2 1 + + ZI– (Z have 3 1 and + + 3 1 . Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 iia,adsm opoercadmrsi hrcesaesmaie nTbeI.The II. Table in summarized are characters of meristic crab and first morphometric some and similar, 1836 codnewt h erdciebhvoro h dls eae of fresh Females to adults. the tolerance of higher behaviour a reproductive the have of with and accordance potential those endotrophic (1998): than higher Schuh a water and show Diesel size, by body obtained in larger those larger with are eggs agree deposited results newly These II). I, (Tables stages of morphology juvenile overall The 1968). Ewald ea ntepseirmri.Teefaue aebe bevdi nysm fthe of sesarmid some other only of serrulate megalopae in of the observed in pairs been found three also have with were features and traits species: furca These similar rudimentary and margin. studied, a posterior individuals with as the setation telson similar or on (3) a with setae stages; maxilla material with zoeal the peduncle of the antennular new endopod the bilobed in of (2) awaiting segment process; spinous first rudimentary (1) I, a Table morphology: zoeal of from remnants antenna, represent it mother. the the omitted of of identification setation therefore the the or of have doubtful confirmation of are uropod We descriptions of and somites, The megalopa abdominal incomplete. maxilliped, the debatable. third maxilliped, of also first maxillule, description is The (2006) I. Table Abrunhosa that in fact the presented in by are larger and is sternum sternite the cheliped between of the differences setation on the setae by of distinguished number genus the easily the be of can species megalopae both for described been previously roberti not seven has with I) and (Table cincinuli; epipod two maxilliped 7. with first 1, endopods and setation pleopod (6) (8) setae with setae; absent; uropods plumose lateral epipod (9) marginal ventrally maxilliped posterior 36–39 second one rostrum with (7) and setae; scaphognathite broad, anterior four maxillar than with two (5) two-segmented three- longer with one; palp peduncle mandibular antennal distal carapace (4) (3) on segments; a absent; setae six endopod with (1) characters antennule flagellum the (2) and (2006b): cleft; with segmented al. medial closely a very et with corresponds deflected It Cuesta Sesarmidae. by family the listed of species material. other identified for accurately and new on based preferably revised, be is ueiewspeiul ecie nyfor only described previously was juvenile first rpoda(usa19;Cet n ne 05.Tehmlg ewe both between homology the The of description 2005). this in inaccuracies Anger presumed of the to development and Due larval doubtful. Cuesta very thus is 1999; characters (Cuesta rae angustipes Armases 1999), 1962), Bookhout and Costlow (see hte hs oalk tutrscnas eecutrdi eaoa olce rmthe from represent collected might megalopae in tested deviations encountered be be to morphological also remains field. can it these structures conditions, zoea-like Since laboratory-rearing these artificial whether 2004). of consequence al. a as et artefacts Guerao (see 1900) ubro etrso h eaoa ecie eenfo oeidvdasapa to appear individuals some from herein described megalopae the of features of number A sternum the of setation the and pereiopods the of segment coxal the on spinulation The of megalopa the of morphology The nweg ftefrtca tg of stage crab first the of Knowledge and aaeam erythrodactyla Parasesarma rtspisonii Aratus .Gea tal. et Guerao G. .ricordi A. .ricordi A. seCet n ne 01,and 2001), Anger and Cuesta (see ersie npripd n ftemglp.Temglpeof megalopae The megalopa. the of 4 and 3 pereiopods on spines bear .ricordi A. .ricordi A. .benedicti A. ssalrta htof that than smaller is H in dad,13)(e anr1968), Warner (see 1837) Edwards, Milne (H. and isladShh(98 nepe hs ifrne in differences these interpret (1998) Schuh and Diesel . ed o nld ti al n ugs hti should it that suggest and I Table in it include not do we , .roberti A. .roberti A. aaeam plicatum Parasesarma Hs,16)(e rewo n ile 1988), Fielder and Greenwood (see 1865) (Hess, rae roberti Armases Armases uha h eaino h eodmaxilliped, second the of setation the as such , hnin than .roberti A. rae rubripes Armases n te eamdcasi iie.The limited. is crabs sesarmid other and , .ricordi A. .roberti A. and olwn h ednyo l larval all of tendency the following , .ricordi A. eieam fasciatum Perisesarma Ltele 86 seSelvakumar (see 1806) (Latreille, .ricordi A. aveof larvae ; , .ricordi A. and .benedicti A. seFgr 3,b) 13a, Figure (see ssmlrt htknown that to similar is .ricordi A. .ricordi A. .roberti A. eam reticulatum Sesarma and , Armases seDı (see yLm and Lima by .rubripes A. (Lanchester, ees their release ac with hatch . Armases . a and ´az Other is Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 ,ashtss d odt;s setae. s, data; no nd, aesthetascs; a, 3 Maxilliped 2 Maxilliped 1 Maxilliped Maxilla Maxillule Antenna w(m 1.17 1.35 Antennula (mm) cw (mm) cl Feature genus the of juveniles first-stage among differences Morphological II. Table A genus crab grapsid the of review A 1992. LG. Abele of those while sea, the into directly larvae ssd rjc uigafedti pnoe yteDush Forschungsgemeinschaft Deutsche the References by sponsored trip Schubart. species D. field Jamaican C. two a to the 1460/3-1) of during (SCHU collection project the with side assisted as Regensburg of the of collection in University facilitated the larvae DE-FG02-97ER12220) from of (grant project of raising Energy a females and of the ovigerous (USA) Department Lafayette and at U.S. crabs Louisiana the of of of al. University the maintenance from et Felder the Darryl Anger with laboratory. helped also Nettelmann (see Uwe currents river by Acknowledgements sea the to transported are 2006). they where from ne .19.Dvlpetlbooyof biology Developmental 1995. K. Anger ´ vrzZ wl .19.Eetsd aslnddyl it or ldsrol avrode larvario desarrollo el sobre dieta la y salinidad la de Efectos 1990. J. Ewald Z, lvarez xpd()89 ,58 ,4 8, 5 4 1, 9 4 5, 8, 1, 2, 12 13 6 6, 5 5, 12, 50 18, 3 4, 9 5, 5 1, 6, 2, 4, 14, 18, 5 6, 10–11 18–20 7 5, 4 nd 3 5, 1, 12, 10 17, 6, 1, 1, 5 5, 5 6–7, 8–9, 11 16, 15 20, 8–9 5, 1, 2, 4 0–2, 41 5 (s) Endopod 20–21 (s) 12 Exopod (s) Endopod (s) Exopod 48 (s) Endopod (s) endite Basial (s) endite Coxal (s) Scaphognathite ailedt s 9 (s) endite Basial oa nie()1 2n 11 17 nd 3 nd 5, 1, 2, 0, nd 12 20 13 3 4, 1, 2, 0, 0, 19–20 14 3 4, 1, (s) 2, endite 0, Coxal (s) endite Basial (s) endite Coxal (s) Flagellum euce()6 ,11,1 d1,1 1 1, 10, nd 1 1, 12, 1 1, 6, (a Exopod (s) Peduncle h ecito fanwgns mtsna otiuin oZooy527:1–60. Zoology to Contributions Smithsonian genus. new a of description the auttv eihtoh flra tgs aieEooyPors eis117:75–81. Series Progress Ecology Marine stages. larval of lecithotrophy Facultative 54:55–60. Marina Scientia ). (, 1853) Edwards, + )0 3–4 0, s) .americanum A. peetpaper) (present rae roberti Armases ¡ ¡ + + + 88 ,230 3 0, 2–3 0, .21.07 1.20 0.02 0.02 510 rae miersii Armases nteGl fMxc.Tba at n oisWeil Tobias and Santl Tobias Mexico. of Gulf the in aveadfrtsaejvnlsof juveniles first-stage and Larvae peetpaper) (present rae ricordi Armases Sesarma .roberti A. ¡ ¡ + + Gasde,aca reigi urtdlrc ol.1. pools. rock supratidal in breeding crab a (Grapsidae), + 7nd 13 ,2n ,4 0, nd 2 0, .2n nd nd nd nd 0.02 0.03 4nd 14 Cutca eaoa rpia)i mrc,with America, in Grapsidae) Decapoda: (Crustacea: ees hi aveit freshwater, into larvae their release rae ricordi Armases wl 1968) Ewald (Dı Armases a and ´az . rae 1837 Armases eam ricordi Sesarma rae rubripes Armases wl 1968) Ewald (Dı a and ´az + ,1 0, (Milne Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 iaJ buhs .20.Tecmlt avldvlpetof development larval complete The 2006. F. Abrunhosa J, Lima usaJ,AgrK 05 avlmrhlg n aiiytlrneo adca rmWs Africa, West from crab land a Garcı of JA, tolerance Cuesta salinity and morphology Larval 2005. K. crab Anger sesarmid JA, the Cuesta of morphology Larval 2001. K. Anger JA, Cuesta Morfologı 1999. JA. of Cuesta development larval The 1962. CG. descriptions. Bookhout of larval Jr, development JD brachyuran larval Costlow complete of The standardization 1960. and CG. Bookhout Accuracy Jr, 1998. JD Costlow GW. with Indies Pohle West DK, the Calazans of PF, decapod Clark terrestrial and freshwater The 1969. Jr. HH Hobbs Jr, FA Chace Gime G, Torres K, Anger 1838 usaJ,ShhM islR cuatC.19.Abeitddvlpetof development Abbreviated 1999. CD. Dı Schubart R, Diesel Sesarmidae M, eleven Schuh of JA, stages zoeal Cuesta first the of Morphology 2006b. CD. Schubart H-C, Liu G, Guerao JA, Cuesta islR cu .19.Efcso aiiyadsavto nlra eeomn ftecrabs the of development larval on (Grapsidae: starvation crabs and Jamaican salinity by of Effects land 1998. of M. invasion Schuh the R, of Diesel reconstruction A 2000. M. Schuh CD, Schubart R, Diesel rewo G ile R 98 avldvlpeto he pce of species three of development Larval 1988. DR. convergent examinations Fielder of on based case JG, systems classification a Greenwood setal and crabs, seta mangrove the of Tree-climbing definition 2005. the Revising CD. 2004. A. Schubart Garm S, Cannicci M, Vannini S, Fratini uroG ne ,Ntemn ,Shbr D 04 opeelra n al ueiedvlpeto the of development juvenile early and larval Complete 2004. CD. Schubart U, Nettelmann K, Anger G, Guerao nl W 92 avlsae fnrhatr tatccas nilsrtdky odn hpa n Hall, and de Chapman London: States populacional key. United Estrutura illustrated the 1994. an of crabs, VGL. Proceedings Atlantic Grapsidae. Kowalczuk northeastern family of the stages of Larval crabs 1992. of RW. Ingle larvae the on Studies 1924. OW. Hyman Herna a ,EadJ.16.Acmaio ftelra eeomn of development larval the of comparison A 1968. JJ. Ewald H, ´az ne gieaJ,TrlAlmaza Toral JL, Aguilera ´ndez eamde,fo h mznrgo,rae ntelbrtr.RvsaBaier eZoologia de Brasileira Revista laboratory. the in reared region, Amazon the 23(2):460–470. from Sesarmidae), laborato rcyr,Gasie) ora fCutca ilg 21:821–838. Biology Crustacean of Journal Grapsoidea). Brachyura, 4:281–294. Crustaceana 33:127–144. Development and United Physiolgy Reproduction Invertebrate Dominica. and of Survey Behaviour 292(5):1–258. Biological Bulletin Freshwater Bredin-Archbold-Smithsonian Museum In: National and Dominica. States Marine to reference stress. special salinity by inhibition versus 39:269–278. odours adult armatum p. 291 Sevilla. de Universidad Seville: 118:203–214. Bulletin Biological laboratory. Cutca rcyr,Gasie)fo h noWs aii,wt umr ffmla larval familial of summary a with Pacific, Indo-West 49(3):151–173. Development the and Reproduction from Invertebrate characters. Grapsoidea) Brachyura, (Crustacea, 79(2):175–196. Crustaceana material. pisonii Aratus eamne,aca htbed nspaitrlrc ol.Junlo rsaenBooy19(1):26–41. Biology Crustacean of Journal pools. rock supralittoral in breeds that crab a Sesarminae), ricordi eamne.Junlo olg 250:141–160. Zoology of Journal Sesarminae). 2:225–248. Supplement ftemuhatsteo ee pce fdcpd.Zooia ora fteLnenScey142:233–252. Society Linnean the of Journal Zoological decapods. of species seven of setae mouthpart the of 7:219–233. Research Ecology Evolutionary evolution. 18(3):423–436. Biology Crustacean roberti A. agoecrab mangrove 21:71–91. Micronesica Australia. Eastern from Grapsidae) aua itr uemPbiain.33p. 363 Publications. Museum History Natural 65(10):1–8. Museum National Me Biodiversidad, la de Direccio Mexico: Marina. rpia)d lad ao,Caioba Farol, do Ilha da Grapsidae) estomato of comparison .Gea tal. et Guerao G. .MleEwrs(rcyr,Gasde erdudrsmlrlbrtr odtos Crustaceana conditions. laboratory similar under reared Grapsidae) (Brachyura, Edwards Milne H. ro[nulse S israin.Crtb Bai) nvriaeFdrld Parana. do Federal Universidade (Brazil): Curitiba dissertation]. MSc [unpublished ´rio pdsydeca y ´podos Bahua rpoda eacnde.Junlo rsaenBooy25(4):640–654. Biology Crustacean of Journal ). Grapsoidea: (Brachyura: Dcpd:Gasde rmJmia ihntso h ilg n clg faut.Junlof Journal adults. of ecology and biology the on notes with Jamaica, from Grapsidae) (Decapoda: aGerr U Rodrı MU, ´a-Guerrero H in dad,13)(eaoa acyr,Gasie)fo laboratory-reared from Grapsoidea) Barchyura, (Decapoda, 1837) Edwards, Milne (H. Parasesarma eieam fasciatum Perisesarma nzL 06 eaopoi fassri ie crab, river sesarmid a of Metamorphosis 2006. L. ´nez pdspr lGlod eio Rı Mexico, de Golfo el para ´podos alra el aii rpia Cutca eaoa rcyr)[otrlthesis]. [doctoral Brachyura) Decapoda, (Crustacea, Grapsidae familia la de larval ´a xc F 3 p. 132 DF. ´xico nGnrld Oceanografı de General ´n and Perisesarma ge ,HnrcxM.20a avlmrhlg ftessri crab, sesarmid the of morphology Larval 2006a. ME. Hendrickx A, ´guez nR,Ri Nun Ruiz RE, ´n Cutca rcyr:Ssria)fo igpr,wt larval a with Singapore, from Sesarmidae) Brachyura: (Crustacea: ,P e eevliet pos-embriona desenvolvimento seu e PR ´, ora fPako eerh26(12):1389–1408. Research of Journal . rae angustipes Armases aNvl Comisio Naval, ´a oJ.19.Epce aaoaa ecrusta de catalogadas Especies 1996. JA. ˜o oBao ap.aPors,Yc Secretarı Yuc. Progreso, a Tamps. Bravo, ´o eam reticulatum Sesarma eaeam rubripes Metasesarma rae angustipes Armases rae benedicti Armases nNcoa aae ooiinoyUso y Conocimiento el para Nacional ´n Dn,15)(eaoa Brachyura: (Decapoda: 1852) (Dana, eam cinereum Sesarma Sesarma rae roberti Armases a erdi h laboratory. the in reared Say rae miersii Armases aa 82(Decapoda, 1852 Dana, Dcpd:Barchyura: (Decapoda: Rtbn and (Rathbun) Rtbn (Decapoda, (Rathbun) rosbcondic sob ´rio Bs)rae nthe in reared (Bosc) rae ricordi Armases tmlto by stimulation : (Grapsidae: Cardisoma Sesarma ¸o e de ˜es ´ceos ade ´a and Downloaded By: [Guerao, Guillermo] At: 06:40 22 October 2007 evkmrS 99 h opeelra eeomn of development larval complete The 1999. S. Selvakumar imranRfC 97 avlnwcln von Larvalentwicklung 1997. C. crab Zimmermann-Ruf tree mangrove the of development larval The 1968. GF. Warner cuatC,Dee .19.Omrgltr aaiisadpntainit ersra aias comparative a habitats: terrestrial into penetration and non- capacities of Osmoregulatory evolution 1998. R. and Diesel , CD, phylogeny, Schubart Molecular 2000. DL. Felder R, Diesel JA, Cuesta CD, Schubart four of stages zoeal First 1998. JA. Cuesta CD, Schubart cu ,Dee .19.Bedn narc ol aveo h eiersra crab semiterrestrial the of larvae pool: rock a in Breeding 1995. R. Diesel M, Schuh imV.19.Pyoeei eainhp mn mrcnseisof species American among relationships Phylogenetic 1996. VH. Niem Dcpd:Gasde eeo nahgl aibeevrnet ora fEprmna aieBiology Marine Experimental of Journal environment. variable 185:109–129. highly Ecology a and in develop Grapsidae) (Decapoda: erctn Sateae hss.Hiebr:Universita Heidelberg: thesis]. [Staatsexamen Betrachtung 2:249–258. Supplement Crustaceana Grapsidae). 47:237–250. (Brachyura, Zoology laboratory of the Bulletin Raffles laboratory. the in reared Grapsidae) Brachyura: td fJmia rb ftegenus the of crabs Jamaican of and study Phylogenetics Molecular Brachyura). (Crustacea: crabs grapsoid 15(2):179–190. American Evolution the within lineages marine Research Plankton of Journal Grapsidae). Brachyura: (Crustacea: Sesarminae 20(1):61–84. American the on remarks and 69:248–330. Crustaceana Grapsidae). aieSine62(3):743–752. Science Marine Armases aveadfrtsaejvnlsof juveniles first-stage and Larvae bl,19 Bahua rpia:Ssria) ultnof Bulletin Sesarminae). Grapsidae: (Brachyura: 1992 Abele, rae miersii Armases Sesarma aaeam plicatum Parasesarma tHiebr.7 p. 70 Heidelberg. ¨t pce rmPnm,wt dniiainkeys identification with Panama, from species rtspisonii Aratus Rtbn19) iemorphologische Eine 1897). (Rathbun Sesarma (subgenus H in-dad)rae in reared Milne-Edwards) (H. Ltele 86 (Decapoda: 1806) (Latreille, Armases rae 1839 Armases Armases ( 5 Sesarma (Brachyura, ) ) miersii