Larval stages, developmental ecology, and distribution of Scyllarus sp. Z (probably Scyllarus aoteanus Powell, 1949) (Decapoda: Scyllaridae)

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Larval stages, developmental ecology,and distribution of Scyllarus sp. Z(probably Scyllarus aoteanus Powell,1949) (Decapoda: Scyllaridae)W. Richard Webber a & John D. Booth ba Museum of New Zealand Te Papa Tongarewa , P. O. Box 467,Wellington, New Zealand E-mail:b National Institute of Water & Atmospheric Research Limited ,P.O. Box 14 901, Kilbirnie, Wellington, New ZealandPublished online: 30 Mar 2010.

To cite this article: W. Richard Webber & John D. Booth (2001) Larval stages, developmentalecology, and distribution of Scyllarus sp. Z (probably Scyllarus aoteanus Powell, 1949) (Decapoda:Scyllaridae), New Zealand Journal of Marine and Freshwater Research, 35:5, 1025-1056, DOI:10.1080/00288330.2001.9517061

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New Zealand Journal of Marine and Freshwater Research, 2001, Vol. 35: 1025-10560028-8330/01/3505-1025 $7.00 © The Royal Society of New Zealand 2001

1025

Larval stages, developmental ecology, and distributionof Scyllarus sp. Z (probably Scyllarus aoteanus Powell, 1949)(Decapoda: Scyllaridae)

W. RICHARD WEBBERMuseum of New Zealand Te Papa TongarewaP. O. Box 467Wellington, New Zealandemail: [email protected]

JOHN D. BOOTHNational Institute of Water & Atmospheric

Research LimitedP.O. Box 14 901, KilbirnieWellington, New Zealand

Abstract Phyllosomata and nistos of Scyllarus sp.Z taken from plankton around and offshore northernNew Zealand and in the Tasman Sea are presumed tobe those of S. aoteanus Powell, 1949, the most com-mon Scyllarus species in this area. The full larval andpostlarval development of this species are described.Three features are characteristic of Scyllarus sp. Zphyllosomata; a dorsal crest on the cephalic shield,dorsal thoracic spines, and a forked telson in middlestages persisting as two prominent spines outreachingthe telson posterior margin in the final stage. Scyllarussp. Z's closest affinities are with a group of Scyllarusspecies whose phyllosomata have forked telsons inmiddle stages and medium to strong posterolateraltelson spines in the final stage. The nisto is low inprofile but distinctive in its armature. Too fewScyllarus sp. nistos have been described to suggest anygroup to which that of Scyllarus sp. Z can be assigned.The spatial and temporal distribution of the larvae andpostlarvae of Scyllarus sp. Z are also described. Al-though early larval stages and postlarvae are foundclose to New Zealand, mid- and late-stage larvae arewidely distributed, some well beyond the known lati-tudinal range of the adults. There is probably an ex-tended egg-bearing and hatching period as early stagelarvae are caught in most months. Variable rates ofdevelopment of the larvae and/or delayed metamor-phosis, are also possible.

M01011Received 8 January 2001; accepted 27 June 2001

Keywords Scyllaridae; Scyllarus; larval morphol-ogy; phyllosoma; nisto; New Zealand

INTRODUCTION

Six species of the slipper lobster family Scyllaridaehave been recorded from New Zealand. Of these,three are in the genus Scyllarus; S. aoteanus Powell,1949, S. mawsoni Hale, 1941, and one further,undescribed species from northern New Zealand (L.Holthuis pers. comm.).

Scyllarus is the most species rich scyllarid genusby far with over 40 species known from temperateand mainly tropical seas (Holthuis 1991). Scyllaruslobsters are typically small (Holthuis 1991) and allthree New Zealand species are less than 100 mmlong. Few specimens of any of the New Zealandspecies have been found in the lobster (adult) phase.Even the best known, S. aoteanus, is rarely collectedand less than 40 adults are held in collections. Farfewer specimens of the other Scyllarus species havebeen collected. Consequently, the biology of NewZealand Scyllarus spp. is poorly known. Between1969 and 1992, however, 878 phyllosoma larvae(phyllosomata) and 49 postlarvae (nistos) of aspecies of Scyllarus were collected within andbeyond the coastal waters of northern New Zealand.They were caught in plankton nets by variousresearch vessels targeting plankton, especially fishlarvae or the phyllosomata of Jasus spp. rocklobsters.

The only New Zealand scyllarid whose larvaehave been described is Ibacus alticrenatus Bate,1888. Its complete development was documented byAtkinson & Boustead (1982) from larvae hatched inthe laboratory and caught in plankton.

At least 57 descriptions of phyllosomata of thegenus Scyllarus have appeared in the literature,although no more than 20 of them have been giventhe names of known species. Most larvae have beendescribed from the plankton as Scyllarus sp. or withan interim alphabetic or numeric name viz. Scyllarussp. A, B, C or I, II, III, etc. (e.g., Prasad & Tampi

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1026 New Zealand Journal of Marine and Freshwater Research, 2001, Vol. 35

1957, 1960a; Johnson 1971a). One or morephyllosoma stages of only eight species have beenlaboratory reared to verify their identity (includingRobertson 1968, 1971; Ito & Lucas 1990).

McWilliam et al. (1995) described the subfinal orfinal phyllosoma stages of 14 Scyllarus species(Scyllarus spp. A-N) from plankton off the west,north, and south-east of Australia. Although nonewere given specific names they discussed theaffinities and possible provenance of their larvae.They examined previous studies of Scyllarus larvaefrom Australian waters and, because there were morespecies represented as larvae than as adults,demonstrated what they called "the fallacy ofreferring unidentified larvae only to those speciesrecorded as adults". Therefore, although we believethe larvae of Scyllarus sp. Z belong to S. aoteanus,we have given it an alphabetic designation becauseother Scyllarus species are present in the same areaand more may be discovered.

Like the phyllosomata, nistos are difficult toidentify although identification can be aided bydirect comparison with juvenile and adult lobsters.Only 14 Scyllarus nistos have been described so farand of these, just two have been further reared to anidentifiable lobster, either from laboratory rearedlarvae or from late stage phyllosomata caught in theplankton.

Baisre (1994) published a review of the place ofphyllosoma larvae in the phylogeny of thePalinuroidea. For Scyllarus phyllosomata, Baisre'sphylogeny agrees with the accepted, lobster basedphylogeny (Holthuis 1991), although it places thegenus closer to Thenus than adult morphology wouldsuggest. Baisre (1994) did not examine species ofScyllarus.

The distribution and ecology of scyllarid larvaehave been less studied than those of palinurids,mainly because scyllarids are less importantcommercially. As with palinurids, early reports onscyllarids came mainly from wide ranging planktonsampling expeditions, and catches were often small(e.g., Gurney 1936; Saisho 1966; Johnson 1971d).More detailed distribution data have come fromlarvae collected as bycatch where other crustaceanshave been the primary focus (e.g., Phillips et al.1981; Rothlisberg et al. 1994). We know of only onestudy of planktonic larval distribution (Yeung &McGowan 1991) in which scyllarid larvae were aprimary focus of sampling.

Three general observations emerge from exami-nation of the literature on scyllarid larval distribu-tion. First, scyllarids differ in their distribution

according to genus, for example Evibacus andScyllarus spp. have generally more inshore distribu-tions than Scyllarides spp. (Johnson 197Id; Baisre1994).

Second, although shallow-water Scyllarus spp.and palinurids both have long-lived phyllosomalarvae which hatch near the coast, those of Scyllarusspp. do not disperse as far offshore as those of thepalinurids. Scyllarus mid- and late-stage phyllo-somata occur both in shelf waters where they can bewidespread (Yeung & McGowan 1991; Rothlisberget al. 1994), and beyond, to 1000 km from the coast(Phillips et al. 1981; McWilliam & Phillips 1983).Their offshore spread, however, is usually much lessthan that of palinurid larvae of coastal species caughtin the same surveys. This was also the situation forScyllarus sp. Z compared with J. edwardsii off thesouth-east of the North Island of New Zealand inFebruary 1998 (Chiswell & Booth 1999).

Third, Scyllarus spp. are often not only thedominant scyllarid species among larvae collectedfrom coastal and near-shelf waters (citations inMcWilliam et al. 1995), but frequently the mostabundant of all phyllosomata in those areas (e.g.,Berry 1974; McWilliam & Phillips 1983;Rothlisberg et al. 1994). Their abundance is due inpart to their prolonged egg-bearing and hatchingperiods and/or repetitive breeding in a single year(Baisre 1994). It is also because they generallyremain close to shore, a result of shorter larval lifeand probably also their behaviour. Currents whichbring about widespread dispersal in palinurids wouldeffect similar transport of Scyllarus larvae if they didnot employ a different vertical migration strategyfrom the palinurids, or some other behaviour tomaintain their place in more coastal water.

Like palinurids, Scyllarus spp. migrate verticallyon a diel cycle. S. bicuspidatus phyllosomata occurnear the surface at night and during the day areconcentrated at 40-80 m in early stages and 80-100 m in mid and late stages (Phillips et al. 1981).Moonlight causes mid- and late-stage larvae to bedeeper at night (by 15-25 m). Eddying adjacent tothe shelf break may have a role in retaining the larvaeof Scyllarus spp. relatively near shore (Johnson1971b; Leeetal. 1992, 1994).

The ecology of the postlarva is not as well known,because nistos are less commonly taken thanphyllosomata. They do appear to swim (e.g., Lyons1970), despite contrary indications (Robertson1968). Those metamorphosing from phyllosomatacollected off eastern Australia were active at nightbut buried in sandy substrates by day (Barnett et al.

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Webber & Booth—Larvae of Scyllarus sp. Z (probably S. aoteanus) 1027

1986). The postlarval phase of at least someScyllarus spp. may be much shorter than that ofpalinurids, which lasts from several days to weeks(Booth & Phillips 1994).

In this paper we describe the morphology of allthe phyllosoma stages and the nisto of Scyllarus sp.Z; describe their spatial and temporal distributions;and give the known distribution of S. aoteanus adultsin New Zealand. The best documented samples ofScyllarus sp. Z were taken with larvae of the rocklobster /. edwardsii. Distribution of the phyllo-somata and pueruli of J. edwardsii have beenextensively studied (Booth & Stewart 1992; Booth1994; Chiswell & Booth 1999) and comparisons ofthe larval distributions of both species are made.

The purpose of this work is to provide a guide toidentification of these larvae in the plankton, tocontribute information to the rather sparse, butgrowing pool of data on Scyllarus speciesdevelopment, to shed light on the recruitmentstrategies of lobster species with phyllosomata, andto contribute to the biology of the New ZealandScyllaridae.

METHODS AND MATERIALS

Phyllosomata and nistos were collected as bycatchin plankton tows which were made at and below thesurface at stations around mainland New Zealandand in the surrounding South Pacific Ocean andTasman Sea during 1969-92. Sampling effort wasnot uniform spatially (more stations were samplednorth and east of New Zealand than elsewhere) orover time. Most specimens came from seasonal nightsamples targeting rock lobster {J. edwardsii) larvaeoff the east coast of the country using a fine-meshed(12 mm cod-end) mid-water trawl (FMMWT) at30 m (see Booth 1994), but others came from muchsmaller, often finer meshed nets towed during thenight or day at various depths. Capture positionswere plotted, and all useful ecological andenvironmental data associated with the larvaeextracted. Except in the 1979-82 and 1987-88FMMWT surveys of rock lobster larvae, no attemptwas made to plot the positions of tows that did notcontain Scyllarus spp. The distribution data for /.edwardsii from these surveys provide a usefulframework against which the larval ecology ofScyllarus sp. Z can be compared. Adult specimensof S. aoteanus were caught by scuba and juvenilesin J. edwardsii puerulus collectors.

Table 1 Key to separate the stages of Scyllarus sp. Z.

Stage Character

1 eyestalk unsegmentedantennule lacking subterminal setaeP3 with exopod budP4 absenttelson spine <Vi length oftelson setae

2 eyestalk segmentedantennule with subterminal setaP3 exopod bud elongate, lacking natatory setaeP4 present as a small budtelson spine >Vi length oftelson setae

3 antennule biramous, with 1 row of subterminal sensorysetae

P3 exopod with natatory setaeP4 bud elongate, lacking setaeP5 present as a small budtelson spine strong, longer than telson setae

4 antennule with 2 rows of subterminal sensory setaeantenna biramousP4 elongate, jointed, setose; exopod lacking natatory

setaetelson forked—spine now continuous with telsonsmall dorsal thoracic spine present above P3 coxauropod present as bud

5 anlennule with 4 rows of subterminal sensory setaeP4 exopod with natatory setaedorsal thoracic spine present above P3 and P4 coxae

6 antennule with about 6 rows of subterminal sensory setaemaxilla 2 without setae, present as unarmed budmaxilliped 1 present as tiny budpleopods present as tiny simple budsuropods with cleft tipsdorsal thoracic spine present above PI, P3. and P4 coxae

7 antennule with about 8 rows of subterminal sensory setaeP5 of 2 segments

8 antennule with about 10 rows of subterminal sensorysetae

slight protuberance on maxilliped 2 indicating exopodslight protuberance on maxilliped 3 indicating exopodpleopods with cleft tipsuropods biramous


pleopods fully biramousdorsal thoracic spine present above PI, P2, P3, and P4


antenna broadmaxilliped 1 with anterior and posterior lobesmaxilliped 2 with exopod budmaxilliped 3 with exopod budP5 of 4 segmentspleopods elongateuropods leaf-shapedgills present as buds

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Measurements were taken from five or morespecimens of each developmental stage (seedescriptions for numbers) and include total length (TL)from the anterior margin of the carapace between theeyes to the tips of the telson spines (phyllosomata) ortelson (nistos); carapace length (CL) from the anteriormargin of the carapace between the eyes to its medianposterior margin; and carapace width (CW), the widthof the carapace at its widest point. Phyllosomata weresorted into 10 stages, based on the frequency of larvaeat a given level of development. The characters usedto distinguish each stage are listed in Table 1; the onlycharacters listed for stage 1 are those which change inthe transition to stage 2; the characters listed for eachsubsequent stage are those which have changedmeasurably from the previous stage. P1-P5 stand forpereopods 1-5.

The material studied is held in the Decapodalarvae section of the Crustacea collection in theMuseum of New Zealand Te Papa Tongarewa.

RESULTS

Developmental stages collectedThe numbers of each stage collected and examinedand the number of specimens caught per tow appearin Table 2 (see also Fig. 14 for distribution by stage).The high number of stage 1 larvae results fromsampling of concentrations of recently hatchedlarvae from relatively few hauls. Comparatively lownumbers of stage 2 were caught, probably becausethey had become dispersed. At all subsequent stagesless than half the number of stage 1 larvae caughtwere taken, presumably mainly because of continueddispersal.

Description of the phyllosoma stages and nisto(Fig. 1-11; Table 3)

Stage ] (Fig. 1A-G)TL: 1.0-1.4 mm (58 specimens measured).Ratio CL:CW = 1:1.13 (five specimens measured).Cephalic shield (Fig. IB) Rather pear-shaped;anterolateral margin quite sharply curved; slightlywider than long; posterior margin with a median

projection. Eyestalk (Fig. IB) Unsegmented,slightly shorter than antennules. Antennule (Fig.1C) Curved laterally distal to mid-point; a slender,plumose seta arising ventrally, distal to mid-point atposition of future inner ramus, similar in reach to theantennule tip; 1 seta and 3 sensory setae apically.Antenna (Fig. 1C) Almost 'A length of antennule,tapering to a point with 2 subterminal setae. Firstmaxilla (Fig. ID) Basal endite with 2 large and 3small setae; coxal endite with 3 setae. Secondmaxilla (Fig. 1F) Tipped with 4 long, plumose setaefanning out from a common base. First maxillipedAbsent. Second maxilliped (Fig. 1G) Of 5 segments(coxa not fully segmented at base), armed asillustrated. Third maxilliped (Fig. 1A,B) Long,reaching forward to mid-point of antennule; of 5segments, coxa with a small, strong ventral spine;long, second segment (incorporating basis-ischium-merus) with an indistinct median constriction at pointwhere exopod bud appears in late stages. Pereopods1-3 (PI-3) (Fig. 1 A,B) Present, no fourth pereopodbud; Pl-2 biramous with setose exopods (each with6 pairs of plumose, natatory setae); P3 with a smallunarmed exopod bud; Pl-3 with a robust coxal spineand a strong subexopodal seta, otherwise armed asillustrated. Abdomen (Fig. IE) Small, hardlytapering, terminating posterolaterally in 2 blunt, bud-like processes, each armed with 3 setae and 1 short,fine spine hardly 'A length of setae.

Stage 2 (Fig. 2A-E)TL: 1.7-2.0 mm (eight specimens measured).Ratio CL:CW = 1:1.10 (five specimens measured).Cephalic shield (Fig. 2B) Anterolateral marginsmore rounded than in stage 1; slightly wider thanlong; median projection slightly raised dorsally;posterior shield margin covering slightly more ofthorax than in stage 1. Eyestalk (Fig. 2B) Withincomplete segmentation at base of eye; slightlylonger than antennules. Antennule (Fig. 2C) A smallseta near base of plumose seta distal to mid-point; 1seta and 4 sensory setae apically; 1 small subterminalseta. Antenna (Fig. 2C) Similar to stage 1; a littlemore than 'A length of antennule. First maxilla (Fig.2D) Similar to stage 1. Second maxilla (Fig. 2D)

Table 2 Scyllarus sp. Z numbers of each developmental stage collected and mean catch per tow. Tows varied induration and gear used.

Stage

Numbers caughtMean catch/tow

1

25414.94

2

181.38

3

606.00

4

343.78

5

502.94

6

883.14

7

742.31

8

731.78

9

1011.60

10

1262.10

All

8784.57

Nisto

492.45

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Fig. 1 Scyllarus sp. Z phyllosoma stage 1. A, ventral view; B, dorsal view; C, antennule and antenna, ventral view; D, first maxillae; E, abdomen, ventral view; 3F, second maxilla; G, second maxilliped. ^

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Fig. 2 Scyllarus sp. Z phyllosoma stage 2. A, ventral view; B, dorsal view; C, antennule and antenna, ventral view; D, mouthparts; E, abdomen and P4 buds,ventral view.

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Similar to stage 1. First maxilliped Absent. Secondmaxilliped (Fig. 2D) Similar to stage 1. Thirdmaxilliped (Fig. 2A,B) Similar to stage 1. Pl-2 (Fig.2A,B) Exopods each with 7 pairs of plumose,natatory setae. P3 (Fig. 2A,B) Exopod bud 4 timeslonger than in stage 1 but unarmed. P4 (Fig. 2A,E)Present as an unsegmented bud almost as long asabdomen. Abdomen (Fig. 2E) Tapering, with bud-like processes tending more laterally than in stage1; spine more robust and more than Vi length of setae.

Stage 3 (Fig. 3A-E)TL: 2.4-3.1 mm (45 specimens measured).Ratio CL:CW = 1:1.11 (five specimens measured).Cephalic shield (Fig. 3B) Similar in shape to stage2 but larger with posterior margin now partlycovering coxa of maxilliped 3. Eyestalk (Fig. 3B)Segmented. Antennule (Fig. 3C) Plumose seta distalto mid-point now supported by a distinct node; 1 setaand 4 sensory setae apically; 1 row of 2 sensory setaesubterminally. Antenna (Fig. 3C) A little more than'/2 length of antennule. First maxilla (Fig. 3D) Coxalendite with a fourth small seta, otherwise similar tostage 2. Second maxilla (Fig. 3D) Similar to stage2. First maxilliped Absent. Second maxilliped(Fig. 3D) Similar to stage 2. Third maxilliped (Fig.3A3) Similar to stage 2. PI (Fig. 3A,B) Exopodwith 9 pairs of plumose, natatory setae. P2 (Fig.3A,B) Exopod with 8-9 pairs of plumose, natatorysetae. P3 (Fig. 3A,B) Exopod with 3-4 pairs ofplumose, natatory setae. P4 (Fig. 3A,B,E) Anelongated bud 3 times as long as abdomen; a small,unarmed exopod bud dorsally; a single terminal and3-4 small subterminal setae. P5 (Fig. 3E) Present assmall, unsegmented bud adjacent to abdomen.Abdomen (Fig. 3E) Now narrower at mid-point,with spines more than twice length of setae, directedposterolaterally to give abdomen a forked tip.

Stage 4 (Fig. 4A-F)

TL: 3.1-4.6 mm (29 specimens measured).Ratio CL:CW = 1:1.11 (five specimens measured).Cephalic shield (Fig. 4B) A little more rounded thanin stage 3; posterior margin now completelycovering coxa of maxilliped 3. Antennule (Fig. 4C)With 2 rows of 2 sensory setae subterminally.Antenna (Fig. 4C) Biramous, outer ramus presentas a stout spine; inner ramus about 4/s length ofantennule, with 3 subterminal setae. First maxilla(Fig. 4D) Basal endite with 2-3 robust, spinulosesetae at tip, and 4 small setae; coxal endite with 5setae. Second maxilla (Fig. 4A) Similar to stage 3.

First maxilliped Absent. Second maxilliped (Fig.4A) Similar to stage 3. Third maxilliped (Fig. 4A)Similar to stage 3. PI (Fig. 4A,B) Exopod with 11pairs of plumose, natatory setae. P2 (Fig. 4A,B)Exopod with 11 pairs of plumose, natatory setae. P3(Fig. 4A,B) Exopod with 8 pairs of plumose,natatory setae. P4 (Fig. 4A,B) Exopod bud unarmedbut 4 times longer than in stage 3, endopodsegmented as in P1-P3. P5 (Fig. 4F) Twice as largeas that of stage 3, unarmed. Thorax (Fig. 4E) A verysmall dorsal thoracic spine above coxa of P3, and ina few specimens above coxa of P4. Abdomen (Fig.4F) With a pair of indistinct uropod buds ventrallyat its narrowest point; tip more strongly forked thanstage 3, with 3 setae near base of each fork.

Stage 5 (Fig. 5A-F)TL: 3.7-7.4 mm (49 specimens measured).Ratio CL:CW = 1:1.11 (five specimens measured).Cephalic shield (Fig. 5B) Similar in shape to stage4. Antennule (Fig. 5C) With 2-segmented pedunclegiving articulated inner and outer rami; 4 rows ofsensory setae subterminally on outer ramus.Antenna (Fig. 5C) Only slightly shorter thanantennule; armed similarly to stage 4. First maxilla(Fig. 5D) Basal endite usually with 3 robust,spinulose setae, otherwise similar to stage 4. Secondmaxilla (Fig. 5D) Similar to stage 4. Firstmaxilliped Absent. Second maxilliped (Fig. 5D)Similar to stage 4. Third maxilliped (Fig. 5A)Similar to stage 4. PI (Fig. 5A,B) Exopod with 12-14 pairs of plumose, natatory setae. P2 (Fig. 5A, B)Exopod with 14 pairs of plumose, natatory setae. P3(Fig. 5A,B) Exopod with 10-11 pairs of plumose,natatory setae. P4 (Fig. 5A,B) Exopod with 5-6 pairsof plumose, natatory setae. P5 (Fig. 5F) An unarmedbud twice length of stage 4. Thorax (Fig. 5E) Adorsal thoracic spine above coxae of P3 and P4consistently. Abdomen (Fig. 5F) Uropod buds moreprominent than stage 4; a single pair of incipientpleopod buds evident in some specimens.

Stage 6 (Fig. 6A-F)TL: 5.0-9.2 mm (79 specimens measured).Ratio CL:CW = 1:1.19 (five specimens measured).Cephalic shield (Fig. 6B) Usually noticeably morerounded than stage 5 anterolaterally andposterolaterally, resulting in a shallow indentationeach side of the posterior median projection; anindistinct, small, longitudinal crest present dorsallyat position of mouthparts; shield now obscuring coxaof PI. Antennule (Fig. 6C) With 3-segmented

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Fig. 4 Scyllams sp. Z phyllosoma stage 4. A, ventral view; B, dorsal view; C, antennule and antenna, ventral view; D, first maxillae; E, left side of thorax, dorsal oview; F, abdomen and P5, ventral view. ij^

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Fig. 5 Scyllarus sp. Z phyllosoma stage 5. A, ventral view; B, dorsal view; C, antennule and antenna, ventral view; D, mouthparts; E, left side of thorax, dorsalview; F, abdomen and P5, ventral view.

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Fig. 6 Scyllarus sp. Z phyllosoma stage 6. A, ventral view; B, dorsal view; C, antennule and antenna, ventral view; D, mouthparts; E, left side of thorax, dorsal 3view; F, abdomen and P5, ventral view. oi

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peduncle; about 6 rows of 1-4 sensory setaesubterminally on outer ramus. Antenna (Fig. 6C)Subequal with antennule; armed with a few smallsetae. First maxilla (Fig. 6D) Similar to stage 5.Second maxilla (Fig. 6D) Reduced to an unarmedbud. First maxilliped (Fig. 6D) Present as a tiny budadjacent to second maxilla (very rudimentary insome specimens). Second maxilliped (Fig. 6D)Similar to stage 5. Third maxilliped (Fig. 6A)Similar to stage 5. PI (Fig. 6A,B) Exopod with 15-16 pairs of plumose, natatory setae. P2 (Fig. 6A,B)Exopod with 16-17 pairs of plumose, natatory setae.P3 (Fig. 6A,B) Exopod with 12-13 pairs of plumose,natatory setae. P4 (Fig. 6A,B) Exopod with 10-11pairs of plumose, natatory setae. P5 (Fig. 6F) Largerthan stage 5, about Vi length of abdomen. Thorax(Fig. 6E) Usually with a dorsal thoracic spine abovecoxa of PI, consistently above coxae of P3 and P4(absent above P2). Abdomen (Fig. 6F) Pleopods 1-4 present as indistinct, simple buds; uropod budswith a shallow cleft.

Stage 7 (Fig. 7A-G)TL: 8.0-11.8 mm (70 specimens measured).Ratio CL:CW = 1:1.31 (five specimens measured).Cephalic shield (Fig. 7B) Similar in shape tostage 6, a few specimens with less roundedantero- and posterolateral margins. Antennule(Fig. 7A,F) About 8 rows of 1-5 sensory setaesubterminally on outer ramus. Antenna (Fig. 7A)Similar to stage 6; 1 or 2 small setae. Firstmaxilla (Fig. 7C) Similar to stage 6. Secondmaxilla (Fig. 7D) Bud slightly larger than stage6; 2-3 tiny setae on anterior margin. Firstmaxilliped (Fig. 7D) Bud larger than stage 6;unarmed and unsegmented. Second maxilliped(Fig. 7D) Similar to stage 6. Third maxilliped(Fig. 7A) Similar to stage 6. PI (Fig. 7A,B)Exopod with 17 pairs of plumose, natatory setae.P2 (Fig. 7A,B) Exopod with 16-17 pairs ofplumose, natatory setae. P3 (Fig. 7A,B) Exopodwith 15-16 pairs of plumose, natatory setae. P4(Fig. 7A,B) Exopod with 13-14 pairs of plumose,natatory setae. P5 (Fig. 7G) With developingcoxal segment; a single lateral seta in somespecimens. Thorax (Fig. 7E) Dorsal thoracicspines present above PI, P3, and P4; spine alsoabove P2 in less than 'A of specimens, alwayssmaller than other dorsal thoracic spines.Abdomen (Fig. 7G) Pleopod buds more obviousand larger than stage 6; uropod cleft deeper thanstage 6.

Stage 8 (Fig. 8A-F)TL: 10.9-16.7 mm (73 specimens measured).Ratio CL:CW = 1:1.28 (eight specimens measured).Cephalic shield (Fig. 8B) Similar in shape to stage7. Antennule (Fig. 8A,E) About 10 rows of 1-5sensory setae subterminally on outer ramus.Antenna (Fig. 8A) Similar to stage 7; 1 or 2 smallsetae subterminally. First maxilla (Fig. 8C) Similarto stage 7. Second maxilla (Fig. 8C) Bud with 2-3tiny setae on anterior margin. First maxilliped (Fig.8C) Bud unarmed but with longer, narrower tip thanstage 7. Second maxilliped (Fig. 8C) A distinct kinkin the basio-ischio-meral segment indicating site offuture exopod bud. Third maxilliped (Fig. 8A,C)Future exopod bud indicated as in second maxilliped.PI (Fig. 8A,B) Exopod with 18-19 pairs of plumose,natatory setae. P2 (Fig. 8A,B) Exopod with 19 pairsof plumose, natatory setae. P3 (Fig. 8A,B) Exopodwith 15-17 pairs of plumose, natatory setae. P4 (Fig.8A,B) Exopod with 14 pairs of plumose, natatorysetae. P5 (Fig. 8F) Similar in length to stage 7; 1lateral and 1 terminal small setae in some specimens.Thorax (Fig. 8D) Dorsal thoracic spines presentabove PI, P3, and P4; !4 of specimens with a spineabove P2. Abdomen (Fig. 8F) Pleopod buds withcleft tips; uropods biramous; a single plumose setaon each fork.

Stage 9 (Fig. 9A-F)TL: 13.4-19.3 mm (101 specimens measured).Ratio CL:CW = 1:1.35 (16 specimens measured).Cephalic shield (Fig. 9B) Longitudinal dorsal crestmore distinct than in earlier stages, anterolateral andposterolateral margins a little more inflated thanstage 8 resulting in a slightly less protruding medianprojection. Antennule (Fig. 9A,E) About 11 rowsof 1-5 sensory setae subterminally on outer ramus.Antenna (Fig. 9A) A little broader based than stage8 with several tiny setae distally. First maxilla (Fig.9C) Similar to stage 8. Second maxilla (Fig. 9C)Bud unarmed but larger and flatter than previousstage with anterior and posterior lobes. Firstmaxilliped (Fig. 9C) Bud larger than in stage 8;unarmed. Second maxilliped (Fig. 9C) Incipientexopod bud larger than stage 8. Third maxilliped(Fig. 9A,C) Incipient exopod bud larger than stage8. PI (Fig. 9A,B) Exopod with 19-20 pairs ofplumose, natatory setae. P2 (Fig. 9A,B) Exopod with19-20 pairs of plumose, natatory setae. P3 (Fig.9A,B) Exopod with 16 pairs of plumose, natatorysetae. P4 (Fig. 9A,B) Exopod with 15 pairs ofplumose, natatory setae. P5 (Fig. 9F) Similar to stage

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aa"a"

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Fig. 7 Scyllarus sp. Z phyllosoma stage 7. A, ventral view; B, dorsal view; C, first maxillae; D, second maxilla, first and second maxillipeds, ventral view; E, leftside of thorax, dorsal view; F, antennule, distal third; G, abdomen and P5, ventral view.

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ooo

pBCL'—<O

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ans

Fig. 8 Scyllarus sp. Z phyllosoma stage 8. A, ventral view; B, dorsal view; C, mouthparts; D, left side of thorax, dorsal view; E, antennule, distal third; F,abdomen and P5, ventral view.

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ftcrcrft

RPCOoo

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Fig. 9 Scyllarus sp. Z phyllosoma stage 9. A, ventral view; B, dorsal view; C, mouthparts; D, left side of thorax, dorsal view; E, antennule, distal third; F,abdomen and P5, ventral view.

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8. Thorax (Fig. 9D) Dorsal thoracic spines presentabove PI, P2 (usually), P3, and P4. Abdomen (Fig.9F) Pleopod buds biramous, longer than stage 8.

Stage 10 (Fig. 10A-G)

TL: 18.6-30.5 mm (126 specimens measured).Ratio CL:CW = 1:1.29 (28 specimens measured).

Cephalic shield (Fig. 10B) Slightly more inflatedthan stage 9 with median projection hardly extend-ing further posterior than posterolateral shield mar-gins; dorsal crest more distinct. Antennule (Fig.10A,F) About 12 rows of 1-5 sensory setaesubterminally on outer ramus. Antenna (Fig. 10A)Considerably broader than stage 9 with several tiny,marginal setae distally. First maxilla (Fig. 10C)Similar to stage 9. Second maxilla (Fig. 10D) Budflat and unarmed but larger than previous stage withmore extended anterior and posterior lobes. Firstmaxilliped (Fig. 10D) Bud noticeably more devel-oped than stage 9 with distinct anterior and poste-rior lobes; unarmed. Second maxilliped (Fig. 10D)With distinct, distally directed exopod bud. Thirdmaxilliped (Fig. 10A,D) With distinct exopod budsimilar to second maxilliped. PI (Fig. 10A,B)Exopod with 20-21 pairs of plumose, natatory se-tae. P2 (Fig. 10A,B) Exopod with 19 pairs ofplumose, natatory setae. P3 (Fig. 10A,B) Exopodwith 19 pairs of plumose, natatory setae. P4 (Fig.10A,B) Exopod with 16-17 pairs of plumose, nata-tory setae. P5 (Fig. 10G) Of 4 segments with a fewtiny setae on distal two segments. Thorax (Fig. 10E)Dorsal thoracic spines present above P1-P4 inclu-sive, stronger than in stage 9 although P2 spine usu-ally smaller than spines above PI, P3, or P4 as inearlier stages. Gills (Fig. 10E) A full complementof lobster-phase gills present as buds at the originsof the third maxilliped and pereopods: maxilliped 3and PI each with one pleurobranch, onearthrobranch, two podobranchs; P2, P3, and P4 eachwith two pleurobranchs, one arthrobranch, twopodobranchs; P5 with one adjacent pleurobranch.Abdomen (Fig. 10G) Pleopods longer than stage 9;uropods now flattened, leaf-like with transparent,minutely spinulose membrane; telson with mem-brane similar to uropods.

Changes in characters during developmentTable 3 lists morphological characters whichundergo appreciable change during developmentfrom the first to final stages. The size ranges (totallength) of stages 1 and 2 are clearly separated whilethe size of the third stage is continuous with that ofstage 4, and those of stages 5-9 overlap considerably

with adjacent stages. The shape of the thoracic shieldusually changes from pear-shaped to rounded in themoult from stages 5-6, but is not consistentlyrounded until stage 8. The cephalic shield dorsalcrest appears in stage 6 and increases a little in sizewith each subsequent stage. While the telsonbecomes forked in stages 3-4 (Fig. 3-4), by stage 8(Fig. 8) its posterior margin is convex and becomesmore so until, in the final stage, the fork is reducedto two strong, posterolateral spines (Fig. 10).

Dorsal thoracic spines appear in a definite order.The spine above P3 appears first in stage 4; a spinemay also appear above P4 at this stage, but only ina minority of specimens. In stage 5, spines areconsistently present above P3 and P4, but neverabove PI. By stage 6 there is almost always a spineabove PI but it is usually smaller and blunter thanin stage 7 at which stage it is consistently present andfull sized. The P2 spine is least consistent in size andpresence, not appearing until stage 7 and then onlyin 25% of the specimens examined. The situation issimilar at stage 8 and it is usually but not alwayspresent in stage 9. In stage 10 the P2 spine is alwayspresent but often quite small and blunt.

Diagnostic characters of the phyllosomataThe phyllosomata are somewhat larger than averagefor Scyllarus species. The cephalic shield is pear-shaped in early stages becoming rounded laterallyin later stages (stages 6-8). In mid stages a dorsalcrest develops on the cephalic shield, dorsal spinesappear on the abdomen in a specific sequence, andstrong spines remain on the telson into the final stage.

Nisto (Fig. 11A-D)TL: 14.4-21.9 mm (10 specimens measured).Ratio CL:CW = 1:1.10 (five specimens measured).Carapace (Fig. 11 A,B) Slightly wider than long,armature blunt, low profile, surfaces mostly smooth;rostrum short, rounded, with a dorsal tubercle; risingto a gastric tooth posterior to rostrum flanked by apair of very small tubercles; a pair of larger but lowprofile cardiac tubercles; posterior cardiac regionwith scattered, indistinct tubercles. Branchial ridgewith a single carina supporting about 9 tuberculateteeth reasonably distinct anteriorly, diminishingposteriorly; an uneven row of 7-9 indistincttubercules lateral to branchial carina; posteriorcorners of carapace roughened but hardlytuberculate; branchial carina terminating anteriorlyat a vaguely defined cervical groove; a ridge of about4 tuberculate teeth running forward from grooveabove inner angle of orbit. Front emarginate, straight

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fta"a"ft

oo

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Pft

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oa-

Fig. 10 Scyllarus sp. Z phyllosoma stage 10. A, ventral view; B, dorsal view; C, first maxillae; D, second maxillae and first to third maxillipeds; E, left side ofthorax, dorsal view; F. antennule, distal third; G, abdomen and P5, ventral view.

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2ft5;NftP_p"

l3P

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73

Fig. 11 Scyllarus sp. Z nisto. A, dorsal view; B, lateral view; C, ventral view; D, right first pleopod, anterior view.

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between antennular notch and orbits; orbits deep,hiding most of eye from lateral view. Anterolateralcarapace spines acute, directed anteriorly; lateralmargins arcuate with uneven serrations; a small,distinct cervical incision posterolateral to orbit;margin posterior to cervical incision with little hintof a gastro-cardiac notch. Posterior marginal grooveshallow with a gentle ridge posterior to it; posteriormargin with a V-shaped median notch. A row ofsetae within front margin of orbit anterior to eye; ascattering of setae medial to each branchial carina;posterior margin with a fringe of small setae.Antennule (Fig. 11A,B) Cylindrical, close-set,between antennae; inner flagellum almost twice aslong, more slender and flexible than outer flagellum,armed with small setae on distal half; outer flagellumshort, cigar-shaped with a row of long setae ventrally(Fig. 1 IB); proximal segment of antennule with adorsal, terminal, forward pointing spine. Antenna(Fig. 11A,B,C) Broad and flat with patches ofsparsely scattered setae dorsally. Anterior margin ofterminal segment (distal squame) with 5 roundedteeth fringed by small setae. Small penultimate (fifth)segment articulated at medial angle of fourthsegment, armed dorsomedially with 3 small, sharpspines; fourth segment (proximal squame) with a lowmedian ridge curving forward to an acute anteriormargin; 2 sharp teeth spaced along outer margin,anteromedial margin with uneven, small, sharp teeth,medial extension projecting up into a short, squareblade with 2 spines (Fig. 1 IB); proximal (third)segment triangular, a low median ridge ending in aforward pointing protuberance on anterior margin,2 small spines on medial face of ridge, lateral marginof segment with two small, sharp spines behindarticulation with fourth segment. Sternum (Fig.11C) Fairly flat, anterior segment (somite IV) with2 small, well separated blunt protuberances;posterolateral corners of posterior segment (somiteVIII) produced into strong, sharp, backward pointingspines (Fig. 1 IB). Pereopods 1-5 (Fig. 11B,C)Short, rather uniform in structure, PI shortest andmost robust, P3 longest, unarmed. Abdomen (Fig.11A, B) Terga of abdominal somites II and III andanterior of IV with low profile median carina.Posterior margin of terga I-IV with a V-shapedmedian notch; of I-V with a fringe of small setae.Terga smooth. Pleura of tergite II angledposterolaterally, serrated; of terga III-V rounded andindistinctly serrated. Hard anterior portion of telsonwith 2 pairs of very faint tubercles; otherwise armedas illustrated. Pleopods (Fig. 11B,C,D) On somitesII-V; biramous, both rami fringed by long, plumose

setae; endopod with rod-like appendix internareaching distal tip of endopod, a cluster of cincinnuliat its tip; basipod with a few plumose setae on thedistomedial margin.

Diagnostic characters of the nistoThe nisto has generally smooth surfaces with fewsetae or spines, and low profile ridges andprotuberances. The carapace has one blunt gastrictooth and a pair of indistinct cardiac tubercles; thereis a single, small cervical incision on the lateralmargin of the carapace; the thoracic sternum bearssharp posterior spines on somite VIII; the abdominalmedian carina of abdominal somites II and III isindistinct and peters out on somite IV; the pleura ofthe second abdominal somite is not pointed butangled, and those of somites III-V are indistinctlyserrated.

Geographic distribution (Fig. 12-16)

The bulk of the larvae and nistos of Scyllarus sp. Zwere taken near the east coast of the North Island,but mid- and late-stage larvae were also widespreadto the north of New Zealand and in the north TasmanSea (Fig. 12, 14). Almost all were caught north ofthe Subtropical Convergence, the majority in areaswith surface water temperatures higher than 16°C(Table 4), a pattern consistent with the knowndistribution of adults (Fig. 13).

The geographic distribution of larvae changedduring development. Early larval stages (stages 1-3) were taken almost exclusively within 80 km ofNew Zealand (Fig. 14A). Mid stages (4-6) weretaken within 80 km of the coast but their distributionalso extended to 500 km from New Zealand (Fig.14B). Late stages (7-10) were widespread frominshore waters to more than 800 km from NewZealand (and a similar distance from Australia) (Fig.14C), but nistos were taken only within 80 km of theshore (Fig. 14D).

All the Scyllarus sp. Z larvae examined here weretaken as bycatch in other (mainly rock lobster larvae)surveys, meaning sampling effort was not uniformspatially or over time, and sampling gear varied.Greatest significance can be attached to larvaldistribution data from widespread, interseasonalsurveys using standard sampling procedures. Fig. 15gives such data from the 1987-88 FMMWT survey.Data from more widespread occasional samplingcarried out between 1979 and 1992, using the samegeneral sampling strategy and resulting in capture ofS. aoteanus, are given in Fig. 16; the full extent ofthis sampling is shown in Booth (1994, fig. 2). These

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Table 3 Scyllarus sp. Z summary of larval development, (ant., anterior; biram., biramous; exop., exopod; end.,endite; endop., endopod; ped., peduncle; post., posterior; pr., pair; P, pereopod; seg., segmented; ses, subexopodalseta; St., setae; sp., spine; sss, subterminal sensory setae; term., terminal; TL, total length; uniram., uniramous; unseg.,unsegmented; vcs, ventral coxal spine; 0, absent.)

Stage

TL range (mm)Cephalic shield

Cephalic shielddorsal crest

EyestalkAntennule

Antenna

Maxilla 1

Maxilla 2

Maxilliped 1

Maxilliped 2exopod

Maxilliped 3exopod

Pereopod 3Pereopod 4

Pereopod 5

Pairs pereopodnatatory setae

PleopodsUropodTelson

Dorsal thoracicspines

Gills

1

1.0-1.4pear-shaped

0

unseg.uniram., unseg.

uniram., unseg.

basal end. 5 St.,coxal end. 3 st.4 term. st.

0

0

0

exop. bud0

0

PI=6:P2=6

00bud-like processes,1 tinv sp., 3 st.0

0

2

1.7-2.0pear-shaped

0

seg.uniram., unseg..1 St.

uniram., unseg.

basal end. 5 St.,coxal end. 3 st.4 term. st.

0

0

0

exop. budshort unseg. bud

0

P1=7;P2=7

00bud-like processes,1 small sp., 3 st.0

0

3

2.4-3.1pear-shaped

0

seg.biram., unseg.,1 row sss,exop. setoseuniram., unseg.

basal end. 5 st.,coxal end. 4 st.4 term. st.

0

0

0

exop. setoselong unseg. bud,exop. budtiny bud

Pl=9:P2=8-9;P3=3-400bud-like processes,1 strong sp., 3 st.0

0

4

3.1^1.6pear-shaped

0

seg.biram., unseg.,2 rows sss,exop. setosebiram., unseg.,narrowbasal end. 6-7 st.,coxal end. 5 st.4 term. st.

0

0

0

exop. setoseseg., exop. bud,vcs, sesunseg. unarmedbudP1 = 11; P2=11;P3=801 pr. simple budsforked,3 st.usually above P3(tiny)

0

5

3.7-7.4pear-shaped

0

seg.biram., 2 seg. ped.,4 rows sss,exop. setosebiram., unseg..narrowbasal end. 7 St.,coxal end. 5 St.4 term. st.

0

0

0

exop. setoseseg., exop. setose.vcs, sesunseg. unarmedbudPI = I2-14;P2=14;P3=IO-ll;P4=5-61 pr. buds (some)1 pr. simple budsforked,3 st.above P3, P4

0

results for the east coast of New Zealand confirm thatlarvae of Scyllarus sp. Z seldom occur south of theSubtropical Convergence.

Vertical distribution (Table 4)

Catch rates of Scyllarus sp. Z larvae were three timeshigher at depths <30 m than at greater depths; incontrast, no nistos were taken at <30 m (Table 4).The highest overall catch rates of larvae were inwaters with a bottom depth of 100-500 m. Nistos,however, were taken mainly in deeper waters.

Temporal distribution (Table 5)

Larvae of all stages, and nistos, were caught bothnight and day (Table 5), but overall larval catch ratesat night were 50% higher. Larvae and nistos weretaken at almost all moon phases.

Early stages (including stage 1 phyllosomata)were taken in most months (Table 5) and, becauseof the sampling strategy, it cannot be concludedthat they were absent at any time of the year. Theseresults suggest a prolonged spawning season (thealternative of prolonged development manifested in

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Webber & Booth—Larvae of Scyllarus sp. Z (probably S. aoteanus)

Table 3 Continued

1045

Stage

TL range (mm)Cephalic shield

Cephalic shielddorsal crest

EyestalkAntennule

Antenna

Maxilla 1

Maxilla 2

Maxilliped 1

Maxilliped 2exopod

Maxilliped 3exopod

Pereopod 3Pereopod 4

Pereopod 5

Pairs pereopodnatatory setae

PleopodsUropodTelson

Dorsal thoracicspines

Gills

6

5.0-9.2rounded orpear-shapedpresent

seg.3 seg. ped.,c. 6 rows sss,exop. setosebiram., unseg.,narrowbasal end. 7 st.,coxal end. 5 st.unarmed bud

small bud

0

0

exop. setoseseg., exop. setose,vcs, sesunseg. bud, 1+ st.

P1=15-16;P2=16-17;P3=12-13;P4=10-ll4 uniram. budsslightly cleft budforked, 2 st.

above P3, P4,usually above PI

0

7

8.0-11.8rounded (rarelypear-shaped)present

seg.3 seg. ped.,c. 8 rows sss,exop. setosebiram., unseg.,narrowbasal end. 7 St.,coxal end. 5 st.bud with 2-3minute st.bud withelongated tipslight rudiment

0

exop. setoseseg., exop. setose,vcs, sesseg., 1+ st.

P1=17;P2=16-17;P3=15-16;P4=13-144 uniram. budscleft budforked, 2 st.

above PI, P3,P4,sometimesabove P20

8

10.9-16.7rounded

present

seg.3 seg. ped.,c. 10 rows sss,exop. setosebiram., unseg.,narrowbasal end. 7 st.,coxal end. 5 st.bud with 2-3minute St.bud withelongated tiprudiment

slight rudiment

exop. setoseseg., exop. setose,vcs, sesseg., 1+ st.

P1=18-19;P2=19;P3=15-17;P4=144 cleft budsbiram., unarmedforked, 1 st.

above PI, P3,P4,sometimesabove P20

9

13.4-19.3rounded

present

seg.3 seg. ped.,c. 11 rows sss,exop. setosebiram., unseg.,not so narrowbasal end. 7 St.,coxal end. 5 st.unarmed flat bud,ant. & post, lobesbud withelongated tiprudiment

rudiment

exop. setoseseg., exop. setose,vcs, sesseg., several st.

Pl=19-20;P2=19-20;P3=16;P4=15biram., unarmedbiram., unarmedforked, 1 st.

above PI, P3,P4,usually above P2

0

10

18.6-30.5rounded

present

seg.3 seg. ped.,c. 12 rows sss,exop. setosebiram., unseg.,broadbasal end. 7 St.,coxal end. 5 st.unarmed flat bud,ant. & post, lobesunarmed bud withant. & post, lobesbud

rud

exop. setoseseg., exop. setose,vcs, sesseg., several st.

Pl=20-21;P2=19;P3=19;P4=16-17biram., unarmedbiram., leaf-shapedforked. St. absent

above PI, P2, P3, P4

Full complement ofsimple buds

Table 4 Spatial distribution of ScyUarus sp. Z phyllosoma larvae (by stage) and nistos. In the first two lines of table:1, <80 km from shore; 2, 81-160 km from shore; 3, 161-800 km from shore; 4, >800 km from shore. All other figuresgive mean numbers of larvae by stage, per tow (irrespective of duration of tow or gear used). (Temp, is surfacetemperature, environmental data not available for all tows. - , no lobsters caught at that stage.)

Stage

Temp.(°C)

Netdepth(m)

Bottomdepth(m)

<16>16<30>30

<100100-500>500

1

_12.451.00

1.00_-

2

__1.431.50

1.002.00

_71

_-1

3

.25

.00

.00

4

_14.571.00

__1.00

5

15.002.00

1.001.603.50

6

21,21.144.19

1.006.573.00

7

1,21,2,31.632.70

_4.131.82

8

1,2.31,2,31.171.94

3.861.38

9

1,2,31,2,3,42.001.39

_2.001.26

10

1,21,2,3.44.001.49

_1.831.48

Nisto

11-2.53

1.002.893.67

All

1.2,31,2,3,49.892.96

1.006.382.41

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Fig. 12 Distribution of all sta-tions at which Scyllarus sp. Z lar-vae and nistos were caught (smallislands are shown larger than theyactually are).

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Table 5 Temporal distribution of Scyllarus sp. Z phyllosoma larvae (by stage) and nistos. In upper part of tablefigures give mean numbers of larvae by stage, per tow (irrespective of duration of tow or gear used). (Night, time fromI h after sunset to I h before sunrise; New, First Q, Full, and Last Q are phases of the moon.) In lower part of table:1, <80 km from shore; 2, 81-160 km from shore; 3, 161-800 km from shore; 4, >800 km from shore. (Temporal datanot available for all tows; -, no lobsters caught at that stage).

Stage

DayNight

NewFirst QFullLastQ

JanFebMarAprMayJunJulAugSepOctMovDec

All

1

6.5029.86

52.0015.311.001.00

1_11-11--1

-

1

2

1.251.50

1.001.382.001.33

1,3_11--1

-_L1

1,3

3

3.006.33

1.008.141.001.00

3_-1--1--1--

1,3

4

3.003.88

1.005.171.001.00_--1I-1----1

1

5

1.003.75

2.134.831.33-

-111,2-3----1

1,2,3

6

1.753.70 i

5.50 '11.253.17 ;1.00

i ;-I :I1,2,31,22----1

1,2,3

7

5.501.56

1.445.451.33.22

2,3

,2,32,2,3,2,32

,2,3

8

1.141.94

2.731.231.831.45

1,2,3232,31,21,2,31,2,31,31--3

1,2,3

9

2.091.50

1.331.991.801.33

1,2,31,22,32,31,21,2,31,2,31,2,3,41,2,3,41,2--

1,2,3,4

10

3.291.67

3.362.351.311.00

1,21,22,32,3,411,2,3111,2,3,41,2,312,3,4

1,2,3,4

Nisto

2.002.47

1.673.102.00---1-1-11-111

1

All

3.485.15

6.445.812.242.03

1,2,31,21,2,31,2,3,41,2,31,2,31,2,31,2,3,41,2,3,41,2,311,2,3,4

1,2,3,4

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— i —165

— I —170 175E Fig. 13 Distribution of Scyllarus

aoteanus lobsters (cross-hatching)and surface oceanic flow patterns:EAC, East Auckland Current;ECC, East Cape Current; NCE,North Cape Eddy; ECE, East CapeEddy; WE, Wairarapa Eddy.

^Snares Is^j I '•*Bounty /Platform /

• - ' ' - • •

these early stages as delayed moulting to subsequentstages is not evident in any other species of scyllaridor palinurid). As larvae developed they occurredover more months until at stage 10 they were takenthroughout the year. Nistos were taken throughoutmost of the year but with no clear seasonal pattern.

DISCUSSION

Larval identityThe developmental stages studied were taken fromplankton north-east and north of New Zealand wheretwo species of Scyllarus are known to occur as adults(S. aoteanus and one unidentified species), andwhere other, undiscovered species may also exist.Very few lobsters of the unidentified species havebeen found, but S. aoteanus adults and juveniles areknown to occur around the north and north-east coastof the North Island (Fig. 13) between Ninety MileBeach and Gisborne, a distribution related to thesouth-east flowing East Auckland Current (EAC)and East Cape Current (ECC). The pattern ofdistribution of Scyllarus sp. Z larvae is consistent

with the distribution of adult S. aoteanus whichindicates that they are probably the same species.

Lobsters of S. mawsoni Hale, 1941 have beenfound off the eastern South Island and as far northas 39°S, off Cape Egmont on the west of the NorthIsland (Museum of New Zealand collection), a dis-tribution too southern for this species to be the sourceof Scyllarus sp. Z larvae. In addition, McWilliam etal. (1995) described the subfinal/final stage of aScyllarus sp. N from plankton off Tasmania, givingstrong circumstantial evidence it is referable to S.mawsoni which also occurs in that region. Scyllarussp. N has not been found in New Zealand waters butdiffers markedly from Scyllarus sp. Z.

We are confident the phyllosomata we havedescribed are of a single species even though nophyllosomata of other Scyllarus species in the areawere collected with them. The 878 larvae examinedfall within a size range, sequence of development,and morphological uniformity in accordance withtheir being one species.

The progressive addition and modification ofdistinguishing features (dorsal crest, dorsal thoracicspines, and strong telson spines) plus the continuityof development of the appendages (Table 3), support

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Fig. 14 Distribution of ScyUarus sp. Z by developmental stage; A, stages 1-3; B, stages 4-6; C, stages 7-10; D,nistos.

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Fig. 15 Stations (open circles)and larval occurrence of Scyllarussp. Z (dark circles) off the eastcoast of New Zealand, 1987-88.For details of gear and samplingstrategies see Booth & Stewart(1992).

the probability that these phyllosomata are of a singlespecies. Significant changes occur in the moult fromthe fifth to sixth stages, most noticeably the morerounded cephalic shield with fuller anterolateral andposterolateral margins. In a few phyllosomata,however, the cephalic shield does not becomerounded until the seventh or even eighth stage. It isalso at stage 6 that the dorsal crest appears andmaxilla 2 changes completely, but appreciablechanges occur with each new stage and stage 6 doesfit the sequence of addition of dorsal thoracic spines(see Tables 1, 3).

Phyllosoma stagingAlthough their presence cannot be ruled out with-out direct observation of hatching, no free swimmingnaupliosoma of Scyllarus sp. Z were identified fromthe plankton and apparently have not been observedin other Scyllarus species. A naupliosoma has, how-ever, been found in various Palinuroidea, e.g., thescyllarids /. alticrenatus Bate, 1888 (see Lesser1974), Scyllarides squammosus (H. Milne Edwards,1837), and S. aequinoctialis (Lund, 1793) (seeRobertson 1969) and the palinurids Jasus edwardsli(Hutton, 1875) and J. verreauxi (H. Milne Edwards,1851).

Ten phyllosoma stages are identified in thedevelopment of Scyllarus sp. Z, which are defined

in Table 1. This indicates there are at least 10 instars,possibly more since 11 identifiable stages have beenobserved to pass through a total of 17 instars, inspecies such as Jasus edwardsii (Kittaka pers.comm). We agree with Johnson (1968) that assigningstages is arbitrary and depends on the characters usedand that these can vary according to the workersinvolved (Phillips & Sastry 1980). Although totallengths of our stages 4-10 overlap with those ofadjacent stages (Table 3), we found more stages thanare usually observed (e.g., seven in S. americanus,reared by Robertson 1968; eight in S. martensii byPhillips & McWilliam 1986 and eight for species A-N by McWilliam et al. 1995; and nine in 5. depressusreared by Robertson 1971).

Phyllosoma affinitiesScyUarus sp. Z is clearly Scyllarus because it is moresimilar in shape and developmental morphology toother species of Scyllarus than it is to any otherscyllarid genus. Although characterised by thecombination of a dorsal crest, forked telson, anddorsal thoracic spines, these characters are notunique to Scyllarus sp. Z. Table 6 lists all Scyllarusspp. reported to date with one or more of thediagnostic characters of Scyllarus sp. Z. Also givenare the main differences from Scyllarus sp. Z wherethere has been enough detail available.

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Fig. 16 Larval occurrence ofScyllarus sp. Z (dark circles) offthe north and east of New Zealand1979-92 (excludes 1987-88 sam-ples of Fig 15). For transects whereno 5. aoteanus were caught seeBooth (1994, fig. 2).

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180

Of the species listed in Table 6 only one, Scyllarussp. b, has been documented as having, simultane-ously, a dorsal crest, telson spines reaching beyondthe uropods in the gilled stage, and dorsal thoracicspines (Phillips et al. 1981; = Scyllarus sp. L ofMe William et al. 1995) as Scyllarus sp. Z does. Thisdoes not necessarily mean, however, that this com-bination is not present in some of the otherphyllosomata; rather we suspect that some of thesecharacters, particularly the dorsal crest, have notbeen observed by some other authors. Furthermore,since numerous species of Scyllarus exist but thephyllosomata of only a few are known with certainty,from widely scattered localities, we cannot establisha definite relationship between Scyllarus sp. Z andany other species.

Scyllarus sp. of Johnson (1979) is notable forbeing the only species besides Scyllarus sp. Zdescribed with "dorsal marginal spines at theinsertions of legs 1, 3 and 4" (in stage 5 in Johnson'slarva; stage 6 in Scyllarus sp. Z). The lack of a spineabove P2 in these species suggests dorsal thoracicspines may appear gradually in a predetermined

sequence through the mid and late stages ofdevelopment as is apparent in Scyllarus sp. Z.

Characters of Scyllarus phyllosomataMost, if not all Scyllarus species phyllosomata haveat least a very small, paired posterolateral or lateraltelson spine in the final stage (its presence or absenceis unclear in a few papers). The majority of speciesdescribed have spines shorter than the telsonposterior margin. A minority, including Scyllarus sp.Z, have a pair of spines outreaching the posteriormargin (and uropods) in the final stage (see Table6). A few species have telson spines in the final stageintermediate in length between the uropods andtelson e.g., 5. kitanoviriosus (Higa & Saisho 1983)and Scyllarus sp. A (Berry 1974).

Eight of the species in Table 6 have dorsalthoracic spines while the remainder may or may nothave these spines as they are not mentioned in de-scriptions as absent or present. This suggests thatlonger telson spines in the final stage are accompa-nied by dorsal thoracic spines although one species,Scyllarus sp. I of Prasad et al. (1975) described as

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03oo

Table 6 Presence in other Scyllarus spp. of one or more of the major distinguishing characters of Scyllarus sp. Z. Lack of detail in some descriptions means themain differences from Scyllarus sp. Z are not always discernible. (Telson fork, telson in which spines outreach the telson posterior margin in the final stage;+, present; x, absent; ?, unknown; P, pereopod; TL, total length.)

Attributed species

5. arctuslS. batei

S. bicuspidatusS. delfini

S. depressusScyllarus sp. A

Scyllarus sp. bScyllarus sp. ?Scyllarus sp.

Scyllarus sp.Scyllarus sp.*

Scyllarus sp.*Scyllarus sp.*

SourceTelson Thorax Dorsal

fork spines crest Location Main discernible differences

Gurney(1936) +Prasadetal. (1975) +

Phillips etal. (1981) +Johnson (1971c) +

Robertson (1971) +Johnson (1971a) +

Phillips etal. (1981) +Johnson(1971b) +Johnson (1977) +

Johnson (1979) +Bate (1888) +

Stephenson(1923) ?Gurney(1936) +

Mediterranean SeaIndian Ocean

W and SE AustraliaJuan Fernandez

W Atlantic OceanSouth China Sea

W and NW AustraliaHawaiiHawaii

SE JapanCape Verde Is

Mediterranean SeaCape Verde Is

Greater TLThoracic spines in stage 8 onwards; further detail notavailableNo dorsal crestShape of cephalic shield; antennae and pleopods moredevelopedThoracic spines above Pl-4 in stages 7-9Thoracic spines above P1-P4 at mid stages; cephalicshield shapeMuch smaller final stage (13.4-14.4 mm TL)Smaller abdomen; thoracic spines above P3 and P4 onlyShape of cephalic shield; smaller TL at samedevelopmental stageShape of cephalic shieldMore accentuated fork in telson; antennae longer thanantennulesMore accentuated fork in telsonMore accentuated fork in telson; antennae longer thanantennules

o

S

VN

oa*f

a

"Originally described under different names—see Johnson (1979) p. 595 for summary.

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stage 6, has a distinctly forked telson but lacks dor-sal thoracic spines. The presence of dorsal thoracicspines does not presuppose a forked telson either.Three such species with dorsal thoracic spines andtelson spines considerably shorter than the uropodsare S. sordidus (Prasad & Tampi 1960b; Sankolli &Shenoy 1973; Tampi & George 1975; Ritz 1977),Scyllarus sp. c (Phillips et al. 1981), and Scyllarussp. B (Johnson 1971a).

Nisto affinitiesThe nisto of Scyllarus sp. Z is clearly distinguishablefrom all other Scyllarus spp. nistos described to date(Table 7). It is bigger than most (at 14.4-21.9 mmTL), and in those species of similar size, there aremorphological differences. Among those of similarsize to Scyllarus sp. Z, Scyllarus sp. of Phillips &MeWilliam (1989) from Hawaiian waters appearsmost similar. As in the case of the larvae, not allnistos are known to species level although seven ofthe 11-12 species in Table 7 are named, five reliably.Of these seven species, just two (S. kitanoviriosusand 5. depressus) have final phyllosoma stages withtelson spines outreaching the posterior margin asdoes Scyllarus sp. Z.

Spatial distributionThe results from this paper (and those from Chiswell& Booth 1999) point to Scyllarus sp. Z havingsimilar distributions to other Scyllarus spp. in thatearly stage larvae are mainly inshore while mid andlate stages become widespread in waters over theshelf and beyond (Phillips et al. 1981; McWilliam& Phillips 1983; Yeung & McGowan 1991;Rothlisberg et al. 1994). Scyllarus sp. Z werefrequently taken well offshore, up to 900 km (oneat 1400 km) north and north-west of the North Island(Fig. 12), a similar distance to the maximum reportedfor S. bicuspidatus off Western Australia (Phillipset al. 1981). The standardised survey of 1987-88,however, indicated a limit to the seaward extensionof Scyllarus sp. Z larvae to the east of East Cape (Fig.15) while those of J. edwardsii were taken at allstations east of East Cape (see also Booth & Stewart1992) and are known to be common right to theLouisville Ridge 1200 km to the east. There was noevidence of Scyllarus sp. Z being associated withmedusae, as reported for Scyllarus spp. off easternAustralia (Barnett et al. 1986).

Stage 1—4 larvae were only taken close to shoreoff the north-east of the North Island (Fig. 14A), theonly region where adults of S. aoteanus are known.Intermediate and late stages were more widespread

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along the east coast of the North Island (Fig. 14B,C),a distribution consistent with current flows (Fig. 13)but probably also influenced by the sampling regime.The East Auckland Current (EAC) is a small,persistent current of subtropical origin flowingsouth-east towards East Cape. Within or adjacent toit are the North Cape Eddy near North Cape and theEast Cape Eddy near East Cape (Roemmich &Sutton 1998), both of which may help to hold long-lived larvae nearshore. Flowing south from EastCape is the East Cape Current, which reflects eastand north-east to form the Wairarapa Eddy(Roemmich & Sutton 1998), and is known to retainpalinurid larvae nearshore (Chiswell & Booth 1999).Larvae hatched in the extreme north of the NorthIsland may also disperse south (see Fig. 14C) on theintermittent, south-moving West Auckland Current(not shown).

Although the east coast of the North Island hasbeen sampled more intensively for phyllosomatathan elsewhere in New Zealand, this is not thoughtto be the only reason most Scyllarus larvae werecaptured there; extensive yet unproductive samplinghas also taken place off the east coast of the SouthIsland and along the west coast (Fig. 15; Boothunpubl. data).

In contrast to Scyllarus sp. Z, few mid- or late-stage phyllosomata of J. edwardsii are taken withinthe shelf break (Lesser 1978; Booth & Stewart 1992;Booth 1994). Thus, larval behaviour probably differsbetween species, which supports the probability thatphyllosomata employ strategies to control theirdispersal. Nistos of Scyllarus sp. Z and pueruli of J.edwardsii, however, have similar inshoredistributions, mainly near or within the shelf break.The capture of nistos only near the shelf suggests thatmany of the Scyllarus larvae carried well away fromshore are lost to the adult population.

Unlike the palinurids (including J. edwardsii), inwhich the latitudinal distribution of larvae seldomextends beyond that of the adults (George & Main1967), some larvae of this scyllarid species appar-ently occur south of, and much further north, thanthe adults. The southernmost record of Scyllarus sp.Z is near Banks Peninsula at 44°S; the northernmost22.5°S, near Tonga, although no larvae were takennorth of the Tropical Convergence. Scyllarus sp. Zlarvae do not, therefore, appear to be entrained byeddies to the same extent as those of J. edwardsii,which probably represents a difference betweenthem in larval behaviour (Chiswell & Booth 1999).Larva] dispersal hundreds of kilometres to the east,north, and west of North Cape may take place via

the seaward extension of the EAC, which flows sea-ward north of East Cape (Fig. 13) (Heath 1985). Lar-vae in the EAC seaward extension may also reachthe south-east trade wind belt and so be carriedtowards Australia. Two drogues released off north-ern New Zealand in the late 1970s which reachedPapua New Guinea, are thought to have been trans-ported by this process (Booth 1983), which offers apossible explanation for the presence of occasionalScyllarus sp. Z larvae in the central and west TasmanSea. There are no records of the presence of S.aoteanus adults in Australia (P. Berents pers. comm.)or its larval stages in east Australian plankton, whichhas been extensively sampled for Scyllarus (andother) larvae (Dakin & Colefax 1940; Me William &Phillips 1983; Barnett 1989; Me William et al. 1995).

The difference between night and day larval catchrates (Table 5) probably reflects larval dispersal togreater depths near dawn, as reported for otherscyllarids (e.g., Phillips et al. 1981) and also forpalinurids (Booth & Phillips 1994). The capture ofnistos in the plankton clearly demonstrates that thepostlarva of this species does swim during at leastpart of its life.

Temporal distributionThe lack of distinct seasonality of early larval catchesis consistent with extended hatching periods and/orvariable rates of development and possibly an abilityto delay metamorphosis to the nisto. In line with this,nistos and juveniles have settled over several monthsduring 1999 on J. edwardsii puerulus collectors atGisborne, on the east coast of the North Island(Booth unpubl. obs.). Extended periods of egg-bearing and hatching probably occur in Scyllarus sp.Z and this has been the interpretation for severalScyllarus spp. (e.g., Phillips etal. 1981; McWilliam& Phillips 1983; Rothlisberg et al. 1994) althoughnot all (Barnett 1989).

CONCLUSIONS

Scyllarus sp. Z is the first species in the genus whosedevelopmental stages have been described from NewZealand. Scyllarus sp. Z is probably S. aoteanusPowell, 1949, the most commonly found of threeNew Zealand species in the genus Scyllarus.

The phyllosomata of Scyllarus sp. Z show closestaffinities with a small group of Scyllarus specieslarvae with forked telsons in middle stages and

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medium to strong posterolateral telson spines in thefinal stage. The nisto's armature is low-profile butdistinctive though the nisto of too few species has beendescribed to suggest a close affinity with any group.

Although both the early larval and the postlarvalstages are found close to New Zealand, mid- and late-stage larvae become very widely distributed, includingareas well beyond the latitudinal range of the adults.

In Scyllarus sp. Z there is probably an extendedbreeding season, possibly also with variable rates ofdevelopment of the larvae, and/or delayedmetamorphosis.

ACKNOWLEDGMENTS

We thank all those whose sampling or help led to thiscollection of Scyllarus larvae. Robert Stewart and KatrinaKrsinich did much of the initial larval sorting and stagingand Trevor Willis drafted the distribution maps, for whichwe are grateful. Thanks also to two anonymous reviewerswho provided encouraging and constructive suggestions.

REFERENCES

Atkinson, J. M.; Boustead, N. C. 1982: The completelarval development of the scyllarid lobster Ibacusalticrenatus Bate, 1888 in New Zealand waters.Crustaceana 42: 275-287.

Baisre, J. A. 1994: Phyllosoma larvae and the phylogenyof Palinuroidea (Crustacea: Decapoda): a review.Australian Journal of Marine and FreshwaterResearch 45: 925-944.

Barnett, B. M. 1989: Final-stage phyllosoma larvae ofScyllarus species (Crustacea: Decapoda:Scyllaridae) from shelf waters of the Great Bar-rier Reef. Invertebrate Taxonomy 3: 123-134.

Barnett, B. M.; Hartwick, R. F.; Milward, N. E. 1986:Descriptions of the nisto stage of Scyllarus demaniHolthuis, two unidentified Scyllarus species, andthe juvenile of Scyllarus martensii Pfeffer (Crus-tacea: Decapoda: Scyllaridae), reared in the labo-ratory; and behavioural observations of the nistosof S. demani, S. martensii and Thenus orientalis(Lund). Australian Journal of Marine and Fresh-water Research 37: 595-608.

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Berry, P. F. 1974: Palinurid and scyllarid lobster larvae ofthe Natal Coast, South Africa. South AfricanAssociation for Marine Biological Research.Oceanographic Research Institute InvestigationalReport 34: 1-44.

Booth, J. 1983: Getting the drift of lobster larvae. Catch'83(8): 12.

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Booth, J. D.; Phillips, B. F. 1994: Early life history ofspiny lobster. Crustaceana 66: 271-294.

Booth, J. D.; Stewart, R. A. 1992: Distribution ofphyllosoma larvae of the red rock lobster Jasusedwardsii off the east coast of New Zealand inrelation to the oceanography. In: Hancock, D. A.ed. Australian Society for Fish Biology Work-shop Larval Biology. Bureau of Rural ResourcesProceedings 15. Canberra, Australian Govern-ment Publishing Service. Pp. 138-148.

Chiswell, S. M.; Booth, J. D. 1999: Rock lobster Jasusedwardsii larval retention by the Wairarapa Eddyoff New Zealand. Marine Ecology Progress Se-ries 183: 227-240.

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George, R. W.; Main, A. R. 1967: The evolution of spinylobsters (Palinuridae): a study of evolution in themarine environment. Evolution 21: 803-820.

Gurney, R. 1936: Larvae of decapod Crustacea: part IIIphyllosoma. Discovery Reports 12: 377-440.

Heath, R. A. 1985: A review of the physical oceanogra-phy of the seas around New Zealand—1982. NewZealand Journal of Marine and Freshwater Re-search 19: 79-124.

Higa, T.; Saisho, T. 1983: Metamorphosis and growth ofthe late-stage phyllosoma of Scyllaruskitanoviriosus Harada (Decapoda, Scyllaridae).Memoirs of the Kagoshima University ResearchCenter for the South Pacific 3: 86-98.

Holthuis L. B. 1991: Marine lobsters of the world. Anannotated and illustrated catalogue of species ofinterest to fisheries known to date. FAO FisheriesSynopsis 125 (also FAO Species Catalogue 13):1-292.

Ito, M.; Lucas, J. S. 1990: The complete larval develop-ment of the scyllarid lobster, Scyllarus demaniHolthuis, 1946 (Decapoda, Scyllaridae), in thelaboratory. Crustaceana 58: 144-167.

Johnson, M. W. 1968: The phyllosoma larvae of scyllaridlobsters in the Gulf of California and off CentralAmerica with special reference to Evibacus prin-ceps (Palinuridea). Crustaceana, Supplement 2,Studies on Decapod Larval Development: 98-116.

Johnson, M. W. 1971a: On palinurid and scyllarid lobsterlarvae and their distribution in the South ChinaSea (Decapoda, Palinuridea). Crustaceana 21:247-282.

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Documents

Larval stages, developmental ecology, and distribution of Scyllarus sp. Z (probably Scyllarus aoteanus Powell, 1949) (Decapoda: Scyllaridae)