ttomtrtfcm »!Oito IfyM Scteitf* »l £ #&. iiw4». UJttf). 2*5-2*1 REPKODl ION AND GROWTH OF THE SMOOTH PEBBLE CRAB PHILYRA LAEVIS (Bdl 18551 AT TWO SITES IN SOUTH AUSTRALIA DURING 1990-91. by stvphrn C. McKjllip* & Ruth v, McKtu lp Summary Ma.n un\ SX\ & M.KiLLtf, R, V, U994) Reproduction and growth ul Ihc- .smooth pebble crab Phihru hf\U (Hell hW>altwosites inSouthAustraliaduring W90-91. 7/wjy. J?, -Sor. S-Ausi.. 11N|4/, 245-25K JQNlfttfctbw, W94. Thegrowthand lite historyolthesmoothpebblecrabWftprt l"<v>.\(Be!!!wnsinterredirnmcollection* rmidt! bi-iwecn July l$9fland December |99] from SultanaPoint ant] Coohowie. South Australia, fi feitfifJ appeared tobescmelp.irousandtoreproducetwice a vc-ar at bothsites. At Sultana huntovieeroiis females wen; common in winter{June-July!andfromlatespringtolatesummer(November February' PswS WStfM* *efc"common from late winterloearly sprintsandalso inearly summer. DistinctcoIhklsornew recruits were found inearly spnne and again ip earlv summer. Recruits gre* i.ipidlv, reaching sexuaJ maturity within lourmonths. The mean sue ofjutllll femalesandtheproportionwhichwereovigeroasliifferedamongstpopulations A tabontforyexperiment showedthat foodavailability affectedgrowthand whetherfemalesbecameoviperous. Wepostulatethatdifferences insizeandoviseryamonesipopulationsofPhdyratuteis may be-causedbydifferencesintheavailabilityoffood, Krv Wciro.s Phthra. pebble crab, life history, intenidul, food, sandrlat Introduction Ihesandllat atCoohowie Nay, 10 km north ofSultana Point, wassampled inthesameway butless frequently The smooth pebblecrab. Philyra laevis(Bell 1855K (November 1990 and January. Maah. April. August IS common onsheltered intertidal sandflafs in southern and December 1991) Western Australia. South Australia, Tasmania and P. laevis is active whilst the BOIKfflAI fc covered by Victoria (Phillips*^/. 19S4> Hale (1976) describes waterduring ebband rising tides, bul remains buried the feeding and courtship behaviour of P faevi\, but in the substratum ul high water or when the sandflai iixilc else is known about this crab. Tasmanun ts completely exposed (Hale 1976). At least 30 and populations hfffi laevis have been reported to host a usually morethan 100 individuals were collected from ucmciteun (genus Caranonemertes) (Bell & Hickman within the interltdal zone where P, laevis occurred 1985) and the trcmatode Microphallus para^rapsi (from low waterto about mid tidal level! between the Smith. 198-MBell I98S). From W90-9I wc sHidjed the timeoflow water and when the sandllat was mundated feeding behaviour of Philyra laevis. finding that by The nstmj tide, At each site at least 160m: of although individuals often fed upon other dead or substratum, consisting of.several haphazardly chosen damaged animals, many were unwilling to teed upon 0.5 metrewide strips from the waters edgelomid tide members of their own species and that Water borne level, werehandsearched toadepthof3.5cm, P. lathis cues from damaged P laevis inhibited feedme was found no tteeper than 3.0 em in the substratum (McKillup & McKillup 1992). During the study, we (McKillup & McKillup unpubl.j. Crabs were frozen collected P laevisover an J8-month period from two and later examined tor sex. whether females were sites 10 kilometres apart in South Australia, and also ovujerous. and the carapace width of all individuals sampled 10other southern Australian sites. These data. was measured to the nearest 005 mm. logelhei with results of a laboratory feeding experiment, areusedloinferthe litehistoryo\'P laevis. Reproductive tottddion and average size oj adid; females at additional sites Methods During January and February 1991, at least 73 Sampli/ix ofPhilyra laevis/row Sultana P>int and Philvra laevis were collected from each of 11 sites m Coohowie SouthAustralia andone inVictoria(Swan Bay within Samples, of Philvra laevis were collected from the Port Philip Bay), and examined tor sex. sue and mtcrlidal sandflat at Sultana Point, lower Yorkc reproductive state as described previously. Peninsula, South Australia <35.0X°S 13744°P.> inJuly I9'M). and then approximately every fourweeks from hihoratnry e.vperitnent on sexual development and September 1990to August V991 and in Decembei 1991. growth Abdominal morphology in theCrustaceacan usually * Department of Zoology, LruverMlv OS Adelaide, South be useo to determine sex: adult males often have a Australia, 5005 Australia, relatively narrow, concavesidedandtapered abdomen. * Present uddre^ Departmeni of Biologs, Central ()ueen.dund Universitv. Rockhampton, Queensland 4-702 whilst leiualcs have a hroader and often circulat Australia abdomen almost as wide as the carapace (Hannoll "Md -. ' McKH.l |JP (& R V M.hlLLUf after 15 min. This method ot feeding prevented the scawaiet in ihe permanent dishes from hecomlne fouled; it was replaced lorinighlly. Crabs wen* inspecledon every working day. individuals which had moulted were examined and sexed as either male lemule or intermediates, .md ,my which had limbed i out were replaced m their dishes. Theexperimentbeganon April 2nd 1991 using new I u-h. rI.ljsAhri;xJtoimiiMrcwlinticmn>srcpdliu>attucgjy=<i!jPuhvie(nxilre(ir/e<mvaol7ta-,1f**rn*olm.uiluelfit irllecCrouiothsofwrioem.IahnedDceocnetimnbueerd 1un9t9i0l alJlacnruaabrsyha19d91mocudlhtpefdl ai leasl once. I982i I 'us was the case lor all i'ltilvnt W'7,, wilh carapaces wider than US mm, hul threemorphological Results type;-, were found in crabs 13,5 mm wide or smaller Ihc two previously described forms, plus individuals $atnp/itix $( Philyra laevis fnun Sultana hunt cW with a convex sided abdomen which, ai its' widest ( rHfhnutr point was approximately hall'thewidthofthecarapace the frfe& structure of Ihc population a(Pli!l\ra hirvi\ if ig I) The last dewiibcel individuals were named al Sultana Point from July 1990 to December 1991 i- 'iTntheermgerdoiwattehso,f 20 females, 20 uncimediates and 20 milalluessirawieerdeinatFilgesasl2.83.5 Imnmmiwdiwdien,taelrl l(eJumlayle1s99:i0t),leaasltl males all of carapace width 10.5 mm or less, wa- 15 mm wide jihI 93% tit tire killer were ovigcrous, Observed in the laboratory Considering thai vw also By early sprine (September 1990), dead males and tumid differences m ilk- si/c and proportion of adull females wen. extremely common and large live lemales which were ovigcrousamongst sites, and thai individuals were not: almost half of the live males differences in adull si/e amongst iunspcedledi collected were smaller than 8 5 mm and appeareti to lUjja)ilies were also reported by Hule (1976), be ,1 dimmer cohort of recent recruits. Only iwo live observation* were made as pail of a manipulative adull females were found; both were morethan 75 mm laboratory experiment designed !o examine the effects wide and ovigcrous. The rcmamdei of the sample ol lutid availability upon growlh and egg production consisted of"intermediates" 7.5 mm W'idc or smaller of Mfilvtit /tun:,. Crabs were placed individually in which were no! present in the July sample. A mm SO \ 50 v _s() deep plastic dishes, caeh tilled ftith lahoraiory experimenl showed lhat intermediateswen 10 ml n\ seawaiti, All individuals were numbered on juvenile lemales (see below). In mid-spring (October the dorsal side or their carapace with non tovu. I990J only onecohortofmales was present, and thesv vvaicrproofink- Six dishes, containingtwo mules, two malcis were large: than the recruits first seen m fhc femalesand two intermediates were placed withineach previous month Thejuvenile females werealso larger of 10 lidded 280 x 3K0 :•: ilO mm deep p|.,-ai.. trays and some relatively small adull females were preseni. containing seawatcr toadepth ol 10 mm. Aeration was bul none whs ovigerotis, In late spring (Novcrnhci not needed The seawaler in the trays maintained a high 1990), uo juvenile females were found; the sample levelnfhumidity whn.h reduced evaporation I'rom the consisted ol aduh females al leasi 9,5 mm wide. 89$ dishes and also provided a marineenvironment forthe of which were ovigcrous, plus males from 4,5 |o Ti,rWaysi rw.iehrsewhkeipcihaCtlTmiruUtmPdtoeumtpedruartiunrgetahnedcuxapleuiriumlcndla.y a1n8o,5lhmermcowhiodret.ofInreelaatrilvyelsyusmmamlelrma(lDecscewmabscipresWeOnti length. together with acohort of luvemle females upto9:3 mm rive traV$ were assigned randomly to a "high food" wide. Dead adultswere again common, but live adults treatment andthe M)crabswithin these were fedevery were also found and 7KV of adult females were working das (from Monday lo Friday), whilst the 30 ovigcrous^ The small nudes and juvenile lemales nabs in the oilier five" (rays were led wcekh. on continued to grow through summer and autumn Wednesday-, .is .. "hiw ffati* treatment. Dil'IcrciU (January lo April 1991) and the number of ovigcrous frequenciesof ieeding providedifferent levelsof food adult females declined (30aV-v. in January. 14% m ilVdlkblllty I" mverlcbajiest.;,^. Calow 1973; Monarly February and 5Vr m March). Prom mid to late- W7S), For each Ieeding the ^ix crabs in a tray were autumn (April and May 1991.1 nojuvenile females were removed placed in sis separate dishesused for feeding found and oniy 1.5% and 2% respectively of aduh only BJld offered odlibitumciushed cockle. Knleh'sio females wereovigcrous, but inearly wintet (June I99l}« \ifihtrhui Lamarck from Sultana Pomi P fut'viv is the percentage ol ovigcrous females had risen lo6Xr, often found feeding on this bivalve in (he field and further increased lw94% by late winter(July 1991) INK K.llupcv- McMlup 1992) All c.at-s leO fot 12 min Theearly spring(August 1991) sample was very similm or less and wen:- returned (o their permanent dishes lo dial of September 1990; dead adult males and ^ P REPRODUCTION AND GROWTH OF PHILYRA L4EVIS 247 30 30 13thJuly 90 19thJan '91 ISfriJune'9 z M c Hi,l4l5 ll ,27b C EiffbikUUfrl1ITTTIT1 * ii' <iii 145 3 '.0 3U ' 30 u 7fhSep,90 23rdFeb '91 23rdJuly'9" CD 15 15 TTTTTTTTtTTTTtTTTT ,ul ii ii,r . , M>'.>,;rrr. — 145 275 145 275 Q 30 0" 6thOct'90 (/> 3 CD Q ll ill. u 3thApr '9 ,9thDec '91 n" 15 (D Q jm- n J , .>'.i;r, 275 14.5 275 145 27.5 30 30 8thDec '90 25thMay '91 -n-M- II Ll ill rl I11nnriiiii 275 145 27.5 Carapace width (mm) Fig. 2. The size structure offemale Philvra laevis collected from Sultana Point, South Australia on 14 occasions between July 1990 and December 1991. Solid bars indicate juveniles; open bars, adults. 248 S. C McKILLUP & R. V. McKILLUP "0 t , 30 Z C 275 275 3 30 u 17thSep'90 CD o ,j L.j i,i 145 275 Q CT J (D Q 27.5 D" CO N" CD Q 27.5 CO CO 30 8thDec '90 in ii gj 275 Carapace (mm) \A/idth Fig. 3. The size structure ofmale Phihra laevis collected from Sultana Point, South Australia on 14 occasions between Julv 1990 and December 1991. REPRODUCTION AND GKOWTH OK FfttlYRA LAEViS 249 (emales were common and the sample oflivc Philxra cohort of juvenile females present in early autumn Uu'Vts consisted entirely ofjuvenile females less than (March 17th 1991) was larger than in January and most K.5 rum wideand males which werealmostall smaller appeared to have moulted to adultsby April 13th 1991, than those collected during the previous month. The In late winter (August 18th 1991) dead adults were December 1991 sample was similar to the one ftom common andjuvenile females plus small males were the previous December, consistingofadult males plus present. Jn early summer (December I9lh 1991) dead adult and some juvenile females, with h$% ofadult adults were also found, very lew live adult females females ovigerous. Dead adults were again common were present and juvenile females were common. OTI the sandtlat at Sultana Point in December 1991 The mean carapace width of adult females was The data for Coobowie were consistent with those always greaterat Sultana Point thanCoobowie (Table from Sultana Point. In late spring (November 20th 1) and a greater proportion of adult females was 1990) only males and adult females were found, but ovigerous at Sultana Point than Coobowie on five in mid summer (January 19th 1991) juvenile lemales occasions when comparison was possible (Table 2). were also present and dead males and adult females Males were not compared since it was impossible to were common. The average carapace width of the distinguish between juveniles and adults. TahlI: I. Comparison ofthemean carapace width (in mm) ofadultfemale Philyra lacvis/row CoobowieandSultana Point tin November ctdicelions were made 17dcivsapart; rth the other5 accnstotis crabs were Ci>liectedfront both sites^>n tin same day, \ — mean, s ~ standard deviation and n — somple size Site Coobowie Sultana Point Dale collected \ November 1990 33 9.23 2.37 125 13.17 2.02 19 Junuurv 1991 17 12,74 1.75 49 13.19 1 73 17 March 1991 52 12.62 1 42 71 13.68 1.72 13 April 1991 85 13.18 f.75 65 14.19 1.77 18 August ISBN 7? 11-82 1.94 none louml 19 December 1991 4 11.72 1.72 -ii 13.50 1.83 Tabu;2. Hie numberofadultfemale Philyra lacvis andthose twigerous in samples atlleetedfrom CoobowieandSultana Pointon6occasnm,\ duringthesamemonth. TheNttvemhtrcolleetnms weremade 17days apart; ontheother5tteeastons crabs were collectedfrom both sites on the same day. Site Coobowie Sultana Point Date collected Collected Ovigerous Collected Ovigerous 3 November 1990 128 19 January 1991 5 49 17 March 1991 52 71 4 13 April 1991 89 (1 65 2 18 August 1991 22 15 19 December 1991 9 22 14 1abu 3. 77u'percentage ofovfgerous adult female Philyra lacvis at 12 sitessampledduring January and February 199J, together with the mean carapace width of adultfemales from each site. Number rff Number and Mean carapace mature lemales percentage width of Date sampled Site collected ovigerous females (mm) 19 January 1991 Edithburgh Bay 9 tUj 12.67 19 January 1991 Sultana Point 49 15 (3D 13.19 (9 January 1991 Coobowie 17 (0) 12.74 !9 January 1991 Hickey's Point 25 •% (8) 13.10 19 January 1991 Staii^bury 33 7 (21) 14 35 20 January 1991 Pmnl Turum 3 14.83 20 January 1991 Rogues Point sn 6 (7) 13.90 29 January 1991 James Well 20 1 en 12.65 29 January 1991 Pine Poml 23 1 (4) 12.33 2 Fehruary 1991 Foul Buy 87 t0) I3.S9 2 February 1991 Stun Bay 70 10 (Mi I3.lt) 13 February 1991 Swan Bay (Vic. ft 56 (85) 17.46 » ?5U \ r Mi KIt I i"- v U V Mr KJI.LUP Rcprofhifiin- conditionandavenj^ Kite rffoftfttlts iff reached sevual maturity and reproduced from late addifiomd tfWJ spring to laic summer (November 1990 lo February Data forthe numbei ofadult females c*'IIceted and 1991). while those first lound in early .umnier the number and percentage whiih weiv OVigafOOg lb) (IXxcmbct I'WO) reached sexual maturity by nud 12 populations .sampled in mid lo lute summer 1991 autumn (April 1991) and reproduced until early winter are in Table 5; The percentage of ovigerous females (June lL)91> Data from Coobowie were consistent with decreased at Suliana Point from January to February, this patternofrecruitment, but suggest that many adult ...onlv samples collectedon ihc 19-20th January 1991 P ldfvi\ at Coobowiedid not reproduce inthe summer woe compared statistically- Nevertheless, the of 1990-01, although daUi were scanty, being onlv ft proportion ovigerous difleicd significantly amongst the November 1990 and January 1991 seven sites sampled on lower Yorkc Peninsula \Z& ' Recruits were found one month after reproducing contingency tabic comparison, di* — 6. Chi squared females were present in laic spring and two momh StttlSti* - 122$ ?<> (MlllSj The :•igMt'u. ..." alter they wvre present in early wuitei. Considering heterogeneity amongst sites was largely due to Ihc that Ihe sandflal was only sampled monthly, that the greaterproportionofovigerous lemalesatSultana Ppmi smallest crab found was 2.8 mm (McKillup & fiable 11 c'uiv-.dering all sites sampled, the highest McKillup unpubl.) and that smaller individuals Were percentage ol ovigerous aduh females (KV* > was at likely tii be overlooked amongst sand grains ami Swan Hay, Vm*hiudimng earlv Fchruaiy (when only detritus, the larval stage o\ R iacvi,^ is likely to bt- of If were omgcjous at Sultana Poinl, see earlier relatively short duration (perhaps only 1-2 weeks) I discussion oi Figure 1). The Swan Ray populationalso Another member of the same genus, this purse crab contained ihc largest Phi/yru (oca. lotind Philvraglobnsa (Fabricius), has a larval stage lastim- Furthermore t-u the seven site-, sampled Horn the II Jays in (he laboratory at an average temperature ot 19 20th )anuary (excluding Point Turton where only 2K°C (Knshnan &. Kannupandi 1990). three lemales wen?collected), ovigerous females were The moult from juvenile to adult form in females lound onlv at sues where the average carapace widlJi appears to coincide with sexual maturity, since only mm .»! Imiales was greater than LV00 (Table 3)- iwo females with juvenile abdomens ofmore than M'H) examined were "vigerous (McKillup & McKillup hthontutn vyu ntneor on wxml dwe\tipw?m unpubl.). A relative Cnon allomelrie1') increase in Male, adult lemale and intermediate** Phdxm Uu>vt\ abdomencomparedtocarapace width during the moult ottered food on live ofseven days per week, moulted lo adulthood is common in braehyurans (Hartnoll SOOncr ttalfl (hose only \h\ once per week ( Fable 4j, 1974), mm At then first moult in the laboratory all males moulted For sites sampled I the l9-20lh January 1991. in males, all lemales lo females and all mlennediaLes a greater proportion of ovigerous adult females was to females, except for thetwosmallest which remained present where the mean carapace width offemales was as the intermediate form unril they moulted again. In relatively large Also, there was a greater proportion all cases the variance ofdays elapsing before moulting ofovigerous lemales at Sultana Point than Coobowie was grealci in the low food treatment, and by on all occasions when comparison was possible. inspection the distributions in ihis treatment w«nc Differences amongst sites were nut caused simply bv .skewed lo the fight. None of the adult females in ihc larger lemales being more likely lo he ovigerous; low lood rrentment, butall in the high food treatment examinationofthedata used tocompileTabic2 showed were ovigerous by July ls>9l thai in November, 60 ofn9 adult females between 8 and P mm wide were ovigerous at .Sultana Point. b..i Discussion none ot 19 collected on the same date and within the Pbiivni fai'vts reproduced twice a year at Sultana same size range was ovigerous al Coobowie Point ami death of most adults during the breeding I-Lirthcnnore- on January lnth KJ9L noneofthe 17aduh moudis suggests Otis species is largely sciiielptirous temates collected fromCoobowie wasovnierous.eve-n kLvniit;. found in caily spring (September 1990 though the carapace widths of these individuals were vI,AHIsKumfldurTdbtJwtuutmutliionJ.u\n\ =clufMvi.tint/iii:icbYetdeem- Ph>l\ia laevfii fir\i muulnxl /" hfah (tttd htwjboit trsunneW^ i "tntt\ Treatmen !.„,J I.-w Fund 1 n • n *^^ Makb il) >A 10 _V78 10 3s> 10 17,10 [•iii;ilei lit 23.5 7 38 LQ 45 20 25.31 h'i.. r0ttJifllr$ itt ' ' (in ill 79 to 3H 10 , KI-PR(HHH_TION AND GROWTH (>!• WWYKt J,Mtl\ Si within the si/e range of the 15 ovigerous females available lo \. paupciuius at Coobowic thanal .Suliana & collected from Sultana Poim. Similarly, on December Point (MeK.Hup Bu,|cr 1979, llJS3: McKillup lOKet) 17th M)l, none ol oadult females fromCoobovvie was P'urthermore. we have wore recently postulated that <>\ Igtja&USj even 'hough mk were within the va;- range intenidal scavengers may generall> be short of food of ovigerous females rrom .Suliana Point. (McKillup & McKillup |W)- All Suuih Aiistialun Resultsofthe laboratory experiment were consistent populations o\ P liwvis sampled contained snialici with food supply allcctmg the frequency oi inoulimp. females on average ihan the population al Swan Ray A sampletfl crabs will contain individualsatdifferent Victoria, .suggesiing that food availability may bi Magesofthe moultcycle In the low lood group, crabs limiting the growth and reproduction of man} aboul to moult would have done so soon after the populations, of this scavenger Numbly, lentales i>| experiment began, hut thosewhich had moulted shortly anotherleucosiid crab. Hbuiia Javvi.s (Bell, I855] wen beforebeingcollected would have lo prepare to moult only found oAigerous during Decemhet.Januaiv. May again under the laboratory conditions of low linni and August at Wellington, New Zealand, but records availability, In contrast, in the high lood group. crab> from other localities suggest the duration Of ibC prepared to moult would do so.and others which had reproductive season vanes amongsl siLes (Wear k recently moulted would have adequate resources Fielder WS5J. available to jjrow and moult again This can explain The hypotheses that differences in food avuilabilav the greater mean, variance and positive skew m the a,rO limiting theduration ofthe reproductive period t$ number o! days before moulting in the low compared Pfuhra Uh'Vi> and that individuals in natural io the high food irealnicni iTahlc 4i. populations ofthis species ate short of h)ud could be We suggest, for the following reasons, that testedb> frequently feedingmarked individuals in the differenees in si/c and theproportionoi ovigerousadult field and comparing their growth and reproductive females amongst sites were caused by differences m output with the icM of the population, the results ol foodavailability Firstly, in (he laboratory, crabs ;n lite these experiments may explain vvny Philxni /t/ev/.v fob high food treatment moulted sooner (and iherelure arelatively short lifespanand two seasonally opposed iirew faster) than those in the low looJ treatment and breeding seasons in South Australia. only females in the high food treatment pnxJuced vgjjv Secondly, adull female P. lacvis al Coobovvie were Acknowledgments consistently .smaller than those at Sultana Point The inlcrtida! scavenger NffltSttiiUSfnmpa'tiitts (Lamarck). WWee wwiisshh ttoo ttihiaannkK AAllaann Btsuulilieeti aanndd KKeciitihn Wwaallkkeerr for aprosohranch snail which occupiesasimilarniehe and u(hieeinrceinicLouuuriaaugcceeummieeinnrttmduduumrriimnniggi titnhhceesssuttiuuuddvyy.,,aunq~dg mAlaann Bottuuileeir is often found feeding with Philyru Uwvi.s, is also MMiicchhaaeell fC'oVaithe'eiss.,, ntH<Vo-inMn pkHd|e*oiilOdtiet*iai Watun\udi MaAlIklue-te•WWeolllks ffoorritlhieoiiir smaller (and hungrier) at Coobovvie than at Sultana critical and eoonnsslliiuucclliivvee ccooimitmuteeniUt.ss oonn aa ddiiaallll 00|| IIhhee Point's and we have postulated there is less food manuscript References tilpiain,/t^*mJ/,n(i|l>S8mXjitAh sNi8ucJlyinllretmhaeiolrillcu-:hiMsitenrryopohral.Ul/u.J/a-,e•'i/<',<)>/\//m<\ M(rtki;arhinti:p.kiiSa.ilCNu(.$w$n}n)usApbaentpwvcimnluur\a-l p^o(l^ycmioprnpihcisvman;ianiiohne Vi<>< R"\. Six lasm- \ll lt<M25. eortvlaled with rcM>uree uvuiluhility anJ jificren^LS in ;fHi\r%si*fMi6aj5&tm-e6mdH8.eiwricnlehkrMmit-cjswSID(OiNOetmheL-p.'riheh[il!cWL5tjaCb.aroPehtif>rtsXii'ToW\MucWinii.t*»«e!rr<oxdfha"rLfmO, cnp.onymxpUNeiatry&io\totifBtvilneu;Oxao-biIUpi.ukl,ptu'UtyepltbAuret.tgttltwul1e.lsegnit/n/nl>Mi;r7<oe/»r.Jip)ph4csOX.lMSnCt2JV2irLri\Oj.-/1u?t>i^^M>lf>r1MJ.ini5\'6ti>|>,l>i6bi-l,ti-l^^'0v, tV [tto&fy The nie:i\!iivnn*iii ol hutljtf'f :is i ('mow. R [JUTSJ Dll 'he re«ul;ilnr> iiiilure ol iudiVidunl rellfcf'ive csiimale ih lood available io pnpii|;aions *n ppivMh: \omrohser\;ilions finm hcshwalersnuiU- .' '/ju<i VtiiXttfiub pmtpvfiutus Ihiii 56. 16*22 Umd 170. 4I5-42H. __j. is. McMttin-. f<. V. (1^921 Inhthilion Ul feedriljj rti MaiVuIWsH!.aMnd \M\:t(yGov'tfrfPftrmekr-ut>,LieSeoauil>lsi 6AfusSttoauttihj..^Ustmlld krtefsip,omnisNileBetlonc.ruIshe(AodI/W'L.4tu)iWsTapleiteefti'iHeeit'<,.lisbiyohmiohtcto(tVeIrbMdb.kUvMCaTtsi4rf>a.ovJHnlrt••n'm IIaSkOiWsCuisiee,oKiidiGi,ry(lsl'e7M4i)iilSaentuitairaoenieirnsyirnowttrahhsp,^r(tenncubectwuetn-an iOncar'elhaKtihoinWt.o 4tIh-e4H,risfe of pred;Uion and vunvulion 27. 131-136. Moriakis !•: tl97H» 5iurvaiion and CtttWth indie gawritpod Hu.ito_'lnoegt(iy1c^s8o":t,l)CAtGLijrWdotewtitiihcie,au.pvpPo.tleIS-l.$l,1luN%nlh-IrwnyoAVibooer^lkvej,..LiruiGrpho(hB>k>j>;),"',TmIkk! I'HfiMt.uv,cirpls^emhorDu,ul.eAs.to<Bil..\Vt,klAAI;NnOrtRs.ii."ioikf.MKHsCT(n\a\C bK,o\r_t.*vK.I.iWrS'M4K)Ft--trCoBouirpoKnNoa1 Kkisiinan, T He KaKM i'A-moi, r. (J9SK7) Larval 0id w*M Viei-)iia, Mvlbournei larval developmem of Hie purse crab Pbifom swbitta Wi ak R 'V'l'HM.tk D R lts>KS>TU- -Ttoaie i^maol tl.ilMMun lijfeHi rcaied in the l.iboi.iiory lf\\lit<bi>>iov<u Mew /.eaJand. iarvae of ihe Urathvura tCnuwiCUU, mi 171-181 l.icujpndai \ /, (hcitttnv /tttfi AAm Wi I M"