Larval Development in the Lutjanid Subfamily Caesioninae (Pisces): the Genera Caesio, Dipterygonotus, Gymnocaesio, and Pterocaesio

BULLETIN OF MARINE SCIENCE. 59(1): 310-369. 1996 CORAL REEF PAPER

LARVAL DEVELOPMENT IN THE LUTJANID SUBFAMILY CAESIONINAE (PISCES): THE GENERA CAESIO, DIPTERYGONOTUS, GYMNOCAESIO, AND PTEROCAESIO

Sally E. Reader and Jeffrey M. Leis

ABSTRACT Larvae of eight species representing all four recognized genera in the lutjanid subfamily Caesioninae were identified by the series method. We describe in detail development of seven of these covering all genera and all the major developmental diversity represented in the material we examined. Larval series were assembled from specimens captured in the Pacific and Indian Oceans by plankton nets, midwater trawls, and night light. A few newly settled specimens captured on reefs were also included. Head spines, fin-spine morphology, meristic values, arrangement of the premaxillary processes, and pigment patterns were used to assemble the series with geographic distributions and adult characters used to confirm the identifications. Caesionine larvae are characterized by elongate dorsal- and pelvic-fin spines, pelvic- fin rays longer than spine, and a spine on the postcleithrum typical of all lutjanid larvae. They are distinguished from other lutjanid larvae by external ornamentation on the fin spines, robust head spines, serrate supraorbital ridges and the presence of only one spine on the upper limb of the outer border of the preoperc1e until quite late in development. Caesio spp. may be distinguished from the other caesionine genera by a single lateral premaxillary process in late larvae, whereas Dipterygonotus, Gymnocaesio and Pterocaesio have two lateral premaxillary processes. Larvae of four of the eight species of Caesio were identified using fin- spine morphology, fin-ray counts, and geographical distribution; they are Caesio cuning, Caesio lunaris, Caesio caerulaurea and Caesio teres and/or xanthonata. Only C. teres and/ or xanthonata is not described in detail. Dipterygonotus balteatus has the unique fin meristics of D Xlm-XIV,9-11 and A m,lO, and also shares with C. cuning the lowest pectoral-ray count with a mode of 18. Robust serrae occur on the fin spines and on the supraorbital ridge. Fin spines are moderate in length. Gymnocaesio gymnoptera has the most slender, elongate body of any caesionine and two serrate leading edges on Dsp2 and Asp2, a characteristic shared only with Pterocaesio chrysozona and C. lunaris. Both G. gymnoptera and D. balteatus typically have 7-8 procurrent caudal rays compared to 9-10 in all other species of the subfamily. Pterocaesio spp. can be distinguished from Caesio spp. by the presence of two lateral premaxillary processes and from D. balteatus by fin-ray counts. G. gymnoptera has a noticeably more slender body shape than the Pterocaesio species described herein. Two of the 10 Pterocaesio species are identified and described: Pterocaesio tile which has a unique fin-ray count, and P. chrysozona with two leading edges on the Dsp2. In addition, we examined a large number of larvae we were able to identify only as Caesio sp. or Pterocaesio sp., some of which constituted apparent monospecific series. None of these has any morphological characters or developmental patterns not found in the seven species described herein.

Lutjanids of the subfamily Caesioninae, commonly called fusiliers, are planktivorous fishes restricted to the warm waters of the Indo-Pacific region. Large semi-pelagic aggregations of caesionines inhabit coral reefs, feeding on zooplankton, in contrast to the benthic carnivores which make up most of the species of other lutjanid subfamilies. Caesionine species constitute a major component of the "wall of mouths" (Hamner et aI., 1988) surrounding Indo-Pacific coral reefs. In one species, Dipterygonotus balteatus, only juveniles are found on reefs, whereas adults move to a pelagic existence in continental shelf waters (Carpenter, 1988). Fusiliers are of minor commercial importance due to their small size and primary occurrence over coral reefs where commercial fishing for small semi- pelagic fishes is difficult or impractical. However, they are significant in multi-

310 READER AND LEIS: CAESIONINE LUTJANID LARVAE 311 species catches and are sold in regional markets for human consumption (Car- penter, 1987, 1988). They are also used as baitfish. Due to their abundance, fusiliers are important forage fishes for large reef and pelagic predators and are also important in the flow of energy and nutrients through food chains of coral reefs, particularly in the transport of these from the pelagic environment to the coral reef (Hamner et al., 1988). While larval development has been described in some lutjanid genera (Leis, 1987; Leis and Lee, 1994; Leis and Bray, 1995) little is known of the early life history of caesionine species. Spawning mode is known for only three caesionine lutjanids: Caesio teres, Caesio caerulaurea and Pterocaesio digramma. All spawn pelagic eggs (Bell and Colin, 1986; Thresher, 1984; Yokoyama et aI., 1994, 1995), as do all other lutjanids for which spawning mode is known. Leis and Rennis (1983) briefly described a lutjanid which has since been identified as Caesio cuning. Kojima (1988) illustrated and described a single 15.4 mm specimen identified as Caesio sp. and Fourmanoir (1976) mentioned a Caesio sp. but provided no information other than meristic data. We cannot identify either of these larvae further. Yokoyama, et aI. (1994, 1995) described the development of reared P. digramma and C. caerulaurea eggs and early preflexion larvae. Overall, lutjanid larvae constitute a relatively small proportion of total fish larvae captured in plankton tows in tropical areas (Leis, 1987). However the majority of lutjanid larvae captured over the continental shelf in the Indo-Pacific are caesionines (Leis, 1987). Therefore, caesioninae larvae can be abundant in plankton tows and are usually much more abundant than members of the other lutjanid subfamilies. The purpose of this paper is to describe the larval development of seven caesionine species, so as to document the variety of larval morphologies and developmental patterns within this subfamily. The seven larval series described here encompass the full morphological diversity of larvae that we observed amongst thousands of specimens we examined. We had access to larvae from the western Indian Ocean, the eastern Indian Ocean, and the western Pacific (from Japan to the IndonesianlPapua New Guinea archipelago, the western Coral Sea and western Tasman Sea). Thus, the geographical coverage of the material available to us included the range of all caesioninae species but two: C. striata and C. suevica which are confined to the Red Sea. R. H. Kuiter (personal communication) has seen what he believes to be an undescribed species very similar to C. cuning on the south coast of Java. None of our material comes from this area. Although we cannot be sure, we believe it is unlikely that any significant characters or ontogenetic patterns within the Caesioninae have been missed in our accounts of seven species. We consider the Caesioninae a subfamily of the Lutjanidae. Caesionine fishes have long been recognized as being related to the lutjanid fishes, but authors differed in their level of recognition of this relationship. Johnson (1980) recognized a separate family, Caesionidae, and most authors have followed his lead (but, see Leis and Rennis, 1983 and Leis, 1987). However, this treatment renders the Lutjanidae paraphyletic (Leis, 1987; Johnson, 1993), and as pointed out by Johnson (1993) "there is definitive evidence in jaw muscles and larval morphology that the lutjanid subfamily Lutjaninae is the sister group of the caesionids." Johnson (1993) recommended, in contrast to his 1980 treatment, that the caesionines be treated as a subfamily of the Lutjanidae, and we support this view. In this paper we more thoroughly document the larval evidence mentioned by Johnson (1993). This evidence will not be discussed in detail here as it will be more comprehensively analyzed in a separate paper. 312 BULLETIN OF MARINE SCIENCE. VOL. 59. NO.2, 1996

This paper constitutes the third in a series on larval development in the lutjanid subfamilies. The first deals with the Etelinae (Leis and Lee, 1994) and the second with the Paradicichthyinae (Leis and Bray, 1995). Papers on the Lutjaninae and Apsilinae are in preparation,

MATERIALS AND METHODS

When establishing identifications, we assumed there are no valid taxa other than those recognized by Carpenter (1987, 1988): these are listed in Table 1. Undescribed species may remain because portions of the Indo-Pacific, with its highly diverse coral-reef fish fauna, remain unexplored or poorly sampled. Geographical distributions of adults were important in establishing identifications of the larvae. We used the distributions in Carpenter (1987, 1988) with additions by Randall et aI. (1990) and Allen and Swainston (1988). Definitions, measurements and abbreviations follow Leis and Trnski (1989) except that eye diameter is of the orbit, not the pigmented portion of the eye. Abbreviations used are: A-anal fin; Asp--anal fin spine; BD-body depth; BL-body length; C-caudal fin; CS-cleithral symphysis; ~orsal fin; Dsp-dorsal fin spine; LL-lateral line; MRL-maximum relative length; PI-pectoral fin; P2- pelvic fin; P2sp--pelvic fin spine; sp--spine. Abbreviations for the spines and ridges of the head are generally based on the bone for which the spine or ridge arises and are adapted from those of Johnson (1984): F-low anterior ridge on the frontal, (no spines); FS-supraorbital ridge of the frontal bone; Pcl-post cleithrum; PI - inner border of the preopercle; PO---outer border of the preopercle (U- upper, and L-Iower, refer to the limbs of the inner and outer preopercular border); (a range given for the PI and PO represent asymmetry, i.e., the counts on the left and right side of the fish differ); PAsp--large spine at the angle of the preopercle; Op---opercle; Sb---subopercle; Io--interopercle; Ptd and Ptv posttemporal dorsal and ventral edge; Scl-supracleithrum; Pe - pterotic ridge, (no spines). Lateral-line scale counts include scales along the route of the lateral line from the posterior border of the head to the posterior end of the hypural plate. In most specimens either the lateral line scales are not fully formed, or they can not be discerned, particularly posteriorly. Forming scales are difficult to count, and scales are frequently missing on the specimens, so many of the counts are necessarily approximate. Following the definition of Leis and Trnski (1989), all specimens described herein are considered larvae, because, although they may have attained full external meristic values, they stilI retain temporary specializations for pelagic life. These specializations are primarily the spination of the head (albeit reduced in larger specimens) and pigment patterns not found in the reef-associated adults or in juveniles except immediately following settlement. Material Examined.-The material utilized in this study was obtained from a wide variety of institutions. Methods of capture, fixation, preservation, storage and time since capture varied widely, and this has had a marked effect on the condition of the larvae and, hence the detail and completeness of the descriptions herein. Shrinkage is especially a problem for preflexion larvae (Leis and Lee, 1994). In some cases the smallest specimens were not the least developed (in terms of fin formation or development of spines on the head). This is presumably due to differential shrinkage among specimens. In such cases we refer to the least developed specimen of a series rather than the smallest specimen. Many specimens are damaged, and the elongate spines of the dorsal and pelvic fins and rays are particularly prone to breakage. The pigment associated with these spines and their intervening mem- branes is frequently missing in part or whole due to damage. This limits the utility of characters based on spine length and fin pigment, which is unfortunate, as these are important in distinguishing species both within and among genera. Most of the specimens we describe here were captured pelagically, usually with plankton nets or mid-water trawls, but also with light-aggregation devices. However, a few larger specimens were associated with reefs when captured. These fishes were recently settled from the pelagic environment onto experimental reefs that were cleared daily. These newly-settled specimens are particularly useful in establishing identifications and in documenting the morphological transitions which take place between the larval and juvenile stages. Newly settled larvae were used in the descriptions of C. caeru/aurea and C. cuning. Where possible, five specimens within each mm increment of body length were examined in detail. Often, particularly for larvae >8 mm, this was not possible due to low numbers of available specimens. However, for some size increments, surplus material was frequently available. In some cases, more than the minimum five per mm increment were examined in detail, but surplus material is not listed in the "Material Examined." Larvae were examined under a dissecting microscope. Precision of measurements varied with the magnification used: this varied from 6 to SOX depending on size of the specimen and the structure being measured. READER AND LEIS: CAESIONINE LUTJANlD LARVAE 313

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Some larvae, especially those which have been long in preservative, have faded pigment. We have tried to be cognizant of this problem in examining specimens and describing ontogenetic changes in pigment patterns, but the reader should be aware that it could impair the accuracy of some descriptions of pigment, especially those based heavily on DANA (ZMUC) material. Institutional abbreviations follow Leviton, et al. (1985). The literature cited section does not include citations based solely on the authors of the genera and species. These are given by Carpenter (1987, 1988).

IDENTIFICA nON No reared material was available, so all identifications are based on the series method. Unfortunately, most of the series have gaps and lack very small larvae «3.5 mm, and with fewer than 4 D spines). Most of the changes that take place within such gaps are obvious, but the exact size at which they occur is unknown. Identification of larvae within these size gaps should not prove a problem. How- ever, larvae smaller than those we describe may prove difficult to identify, and further work is required in this area. Until larvae of all species are identified there remains the possibility that some of the putative series contain more than one species. It is with this possibility in mind that we draw the reader's attention to variation that may indicate the series are not monospecific. Lutjanidae.-Larvae were identified as lutjanids through the characters listed by Leis and Rennis (1983) and Leis (1987), including 24 myomeres; very long dorsal-fin spines (particularly the second spine) and pelvic-fin spine, which are smooth to coarsely serrate, but without recurved spinelets; longest pelvic fin-ray as long or longer than pelvic fin-spine; strong head spination, absence of serrae on preopercular spines (except in some species with very fine serrae on the preopercular angle spine); no supraoccipital crest or spines; a small spine on the postcleithrum; and a series of ventral melanophores on the tail at least in preftexion larvae. Caesioninae (Table l).-The fin counts of D X,15, A m,12 are diagnostic for most members of the subfamily Caesioninae (except in C. cuning-A 111,11;C. lunaris-D X,14, A m,l1; P. tile-D XII,20-21, A m,13; D. balteatus-D XIV,IO, A, m,lO) as this combination of counts is not found in other lutjanids. In larger caesionine specimens, the separate ascending process of the premaxillae is unique among lutjanids, and the PI-ray counts of 17-24 in all species except D. balteatus (which has 16-19) will distinguish caesionine larvae from most other lutjanids only a few which have 18 or 19 PI rays and none of which have more than 19 rays (Anderson, 1987; Carpenter, 1987). Caesionine larvae are further characterized by external ornamentation of fin spines in the form of serrae on the trailing and leading edges of the spines (Fig. 1). All species have: two trailing edges on all fin spines and two leading edges on the P2 spines (some species have two leading edges, or a double row of serrae, on Dsp2 and Asp2); late development of scales; only one spine on outer border of upper limb of preoperc1e until after 10 mm; late development of Sb spination; and serrae on the FS. Larvae of Pterocaesio and Caesio cannot be distinguished at the generic level. Neither apparently derived larval characters nor general similarity allow larvae to be placed into groups corresponding to these two genera as they are presently recognized (Carpenter, 1987, 1988). This is not surprising, as both genera as presently conceived are paraphyletic (Carpenter, 1990). Larvae of many of the caesionine species are extremely similar. The main characters used to distinguish among species are fin-spine structure, meristic values, some minor pigment characters, body shape (in the case of G. gymnoptera) READER AND LEIS: CAESIONINE LUTJANID LARVAE 315

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Figure 1. Dorsal-fin spine 2 and pelvic-fin spine serrations. Note: A and B are typical of C. caerulaurea, C. lunaris, D. balteatus, G. gymnoptera and P. chrysozona; C and D typical of P. tile and C. cuning. Scale bar on A = 0.16 mm (approximate) and applies for all photographs A. P. chrysozona (7.8 mm) dorsal-fin spine 2. B. P. chrysozona (7.8 mm) pelvic-fin spine. C. C. cuning (7.3 mm) dorsal- fin spine 2. D. C. cuning (7.3 rom) pelvic-fin spine. and geographical distribution. While we were successful in identifying seven species using these characters, not all species could be distinguished and we were left with a number of multi-species groups. Most of the individuals in these multi- specific groups resemble P. chrysozona. CAES/O. A single lateral process on the premaxillae in all Caesio spp. serves to distinguish larger larvae of them from all other Caesionines (Carpenter, 1988: fig. 5). In addition, 9-10 procurrent caudal rays and adult teeth on the premaxillae in the larger larvae further eliminate D. balteatus and G. gymnoptera which have 7-8 procurrent caudal rays and lack adult teeth on the premaxillae in larger larvae. CAES/O CAERULAUREA.A series between 3.8 mm and 28 mm from the Bismarck Sea, East China Sea, western Coral Sea, western Tasman Sea, and northwest continental shelf of Australia (eastern Indian Ocean) was assembled. Specimens between 11 mm and 17 mm are unknown. This series was linked through: fine to moderate serrae on the fin spines; a single leading edge on all D and A fin spines; head spines moderately robust and smooth; three melanophores on the ventral midline of the tail in preftexion larvae from posterior edge of anal anlage to the fin fold near the caudal anlage; a melanophore on the anterior edge of CS to 6 mm; and no melanophore on the preanal median finfold (Figs. 2-4). Based on distribution this series could be C. lunaris, C. caerulaurea, C. teres or C. cuning, the only Caesio species recorded from the Great Barrier Reef and Coral Sea. Larger fish of this series have D X,15 and A III,12, which eliminates C. cuning and C. lunaris, both of which have only 11 anal-fin rays. Fish of this series have 57-65 LL scales whereas C. teres has a LL scale count of 51-61, mode 55, (Table 1) thus confirming an identification of C. caerulaurea. READER AND LEIS: CAESIONINE LUTJANID LARVAE 317

CAESIO CUN/NG. A series of larvae from 2 mm to 23 mm with a gap between 12 mm and 16 mm was assembled from the Gulf of Thailand, Solomon Sea, Bismarck Sea, western Coral Sea, and western Tasman Sea. The series is linked by: comb-like serrae on trailing edges of all fin spines (Fig. lC, D); all D and A fin-spines with a single leading edge; moderate to robust, smooth head-spines; three melanophores on the ventral tail in preftexion larvae from posterior anal anlage to fin fold near caudal anlage; one melanophore just anterior to the CS to 6 mm (Figs. 8-10). Larger larvae can be distinguished by a long Dsp2 (Fig. 6) and the anal fin count of 111,11;the latter is found only in C. cuning and C. lunaris amongst Caesio spp. However C. lunaris can be eliminated because it has only 14 D rays (rarely 13 or 15) compared to 15 D rays (rarely 14 or 16) in C. cuning, (Table 1). The D and A fin counts and the comb-like serrae of the fin- spine trailing edges serve to link the series across the gap from 12 mm to 16 mm. CAESIO LUNAR/S. A series from 4 mm to 9 mm was assembled from the East China Sea, Japan Sea, Philippine Sea, Andaman Sea, Banda Sea, western Coral Sea and northwest continental shelf of Australia (eastern Indian Ocean). The series is linked by: moderate to course serrae (not comb-like) on the fin spines; two serrate leading edges on Dsp2 in all specimens and on Dsp3 from 5 mm; moderately robust head spines with serrate edges on the PAsp in all specimens; two melanophores on the ventral tail (notochord pigment unknown due to lack of preftexion larvae), one melanophore at the base of lower mid C ray; a melanophore on the anterior edge of CS to 6 mm; preanal median finfold pigment absent in our smallest specimen (4.5 mm) (Figs. 11, 12). Among all caesionines the fin count of D X,14 (rarely 13 or 15) and A III,11 (rarely 10) is found only in C. lunaris and occasionally in C. cuning (Table 1). Larvae of the two are distinguished by the presence of comb-like serrae on the fin spines of only C. cuning (Figs. 8-10). CAESIO TERES AND/OR XANTHONATA.A series from 4.0 mm to 27.5 mm was assembled from the western Coral Sea, the northern Solomon Sea, east Indian Ocean to the southwest of Sumatra, and the Andaman Sea. The series is linked by: fine serrae on the fin spines; single leading edge on all D and A spines; moderately robust, smooth head spines; two ventral melanophores on the tail (notochord pigment unknown due to lack of preftexion larvae), one melanophore at the base of lower C ray; melanophore on anterior edge of CS in smallest specimen only; pigment absent on preanal median finfold. With LL count of 52-56, of the Caesio spp, the assembled series could only be C. lunaris, C. suevica, C. xanthonata or C. teres. Two of these species can be eliminated: C. suevica is limited to the Red Sea and C. lunaris has ]4 (rarely 13 or 15) D rays as compared to 15 (rarely ]4 or 16) (Table 1). We are unable to distinguish between C. teres and C. xanthonata or to be certain that the series is monospecific. This series does not differ signif- icantly in development from the other caesionines covered in this paper and is therefore not illustrated or described in detail. This series most resembles P. chrysozona and D. balteatus amongst those we describe and can be distinguished by very fine spine serrae, finely spotted Dsp2 chevron pigment and a PI count of 20-23 compared to 18-20 for P. chrysozona and 17-19 for D. balteatus (Table 1). D/PTERYGONOTUS (MONOTYPIC). Dipterygonotus balteatus: A series from 2 mm to 21 mm largely from the North West continental shelf of Australia (eastern Indian Ocean) but with some specimens from the western Coral Sea, western Tasman Sea, and Bismarck Sea was assembled and linked together through: moderate serrae on the fin spines; a single leading edge on all fin spines except P2 318 BULLETIN OF MARINE SCIENCE. VOL. 59. NO.2. 1996 throughout development; early forming robust, smooth, head-spines; initiaIly four melanophores on the ventral midline of the tail in preflexion larvae from just posterior to anus to caudal anlage; a melanophore each anterior and posterior to the CS to 3 mm, then only anterior to the CS to 5 mm; absence of melanophores on the preanal median finfold (Figs. 13, 14). In larger specimens unique fin counts of D XIV,1O (rarely XII-XV, 8, 9 or 11), A III,1O (rarely 9 or 11), PI 18 (16- 19), 7-8 procurrent C rays (Table I) and the absence of adult teeth on the premaxillae in larger larvae confirm the identification of Dipterygonotus balteatus (the latter two characters are shared only with G. gymnoptera). GYMNOCAESIO(MONOTYPIC). Gymnocaesio gymnoptera: A series from 7 mm to 30 mm from the Eastern Indian Ocean was assembled and linked together through moderate serrae on the fin spines; the presence of two serrate leading edges on Dsp2 from 7 mm, and Asp2 from II mm; moderately robust, smooth head-spines. In larger specimens 7-8 procurrent caudal rays, no adult teeth on the premaxillae (these two characters shared only with D. balteatus), and the very slender body enabled identification as Gymnocaesio gymnoptera (Figs. 20, 21). We are unable to identify preflexion larvae of G. gymnoptera and we are not certain that the specimens between 7 mm and 10 mm are G. gymnoptera. This uncertainty arises because the body has not yet become particularly slender and development of the procurrent C rays is incomplete. However, these smaller specimens came from the same samples as the larger G. gymnoptera, and because they share most of the elements of the pigment pattern of the larger larvae, it is likely these smaller larvae are G. gymnoptera. PTEROCAESIO.Caesionine larvae that have two lateral processes on the premaxillae, and that lack the meristic characters of Dipterygonotus and Gymnocaesio were identifiable as Pterocaesio. PTEROCAESIOCHRYSOZONA.A series of specimens from 3 mm to 13 mm and one each of 17 rum and 18 mm was assembled, primarily of specimens from the north west continental shelf of Australia (eastern Indian Ocean), with additional material from the Andaman Sea, Philippine Sea, western Tasman Sea and western Coral Sea. The series is linked through: medium to course serrae on the fin spines (Fig. 22A, B); two serrate leading edges on Dsp2 throughout development; moderately robust head spines which are smooth to 10 mm (Fig 23), after which PAsp has small serrae (larvae >17 mm have small serrae on the spines adjacent to the PAsp), (Fig. 24B, C); initially four melanophores on the ventral midline of the tail in preflexion larvae from just posterior to the anus to caudal anlage, (Fig. 22A, B); one melanophore anterior to the CS up to 9 mm, and an additional melanophore posterior to the CS in larvae <4 mm; a melanophore on the preanal finfold to 4 mm. Larger specimens are linked by 9-10 procurrent caudal rays and adult teeth on the premaxillae, (eliminates Gymnocaesio and Dipterygonotus). Two lateral processes on the premaxillae from 6 mm serve to distinguish Ptero- caesio from Caesio which has only one lateral process. The larvae are linked across the gap in the series from 13 mm to 17 mm by a PI count of 17-20 (Table 1), the pigment pattern, two leading edges on Dsp2 and the serrate PAsp. Much of our material of this species is from the extreme eastern Indian Ocean over the continental shelf of northwest Australia and the western Pacific between Japan and northern Taiwan: this distribution eliminates all Pterocaesio spp other than P. chrysozona and P. digramma; the latter can be eliminated by the count of PI 20-23 (Table 1). This confirms an identification of P. chrysozona. PTEROCAESIOTILE. A series from 3 mm to 16 mm with a gap between 10 mm and 16 mm was assembled from the East China Sea, Philippine Sea, western Coral Sea, Andaman Sea and Mozambique Channel. The series was linked READER AND LEIS: CAESIONINE LUTJANID LARVAE 319 through: comb-like serrae on trailing edges of fin spines from 3 mm; only Dspl- 4 with serrae on trailing edges, remaining Dsp smooth throughout development; single leading edges on all Dsp and Asp; moderately robust head spines with serrae present on PAsp after 5 mm (Fig. 29); three ventral melanophores on the midline of the tail in preflexion larvae from the posterior anal anlage to the fin fold near the caudal anlage; one melanophore on the anterior side of the CS to 5 mm; one melanophore on the dorsal notochord tip and early appearance of urostyle pigment (Fig. 28); no pigment on the preanal median finfold; and late development and low number of FS ridge serrae. The following serve to link the series across the gap: late ossification of Asp3; fin-ray counts of D X-XII, 19- 22, A III,13; 9-10 procurrent dorsal rays (Table 1); and comb-like serrae on trailing edges of fin spines (except Dsp5-12). The fin counts are unique among caesionines (Table 1) and confirm an identification of P. tile.

DESCRIPTION OF DEVELOPMENT The shared developmental features of caesionine species are given here in a general description. Differences among genera and species are covered in the sections on individual taxa. Morphology.-Caesionine lutjanid larvae are compressed and of moderate body depth with 24 myomeres. The youngest larvae are moderately elongate with the steep profile of the forehead and deep anterior myomeres of the trunk giving a slightly hunch-back appearance. Initially the body is deeper at the pelvic base than at the anus. The gut which is in the process of coiling in the smallest spe~c- imens, deepens, taking on a more triangular shape. There is a small gap between the anus and the origin of the anal fin which disappears shortly after flexion is complete. After flexion the larvae become more elongate with growth and the profile less steep. In the available specimens, the gas bladder is generally large but not conspicuous because of its position deep within the trunk musculature and not visible in all specimens. Some specimens have exceptionally expanded gas bladders, undoubtedly due to rapid ascent to the surface in the net. The head is of moderate size and compressed. The snout length is about the same as the eye diameter. The snout is initially triangular but as the head profile becomes less steep, the snout becomes more rounded. The nasal pit bridges over to form two nostrils at 7-9 mm. The mouth is large and moderately oblique reaching at least the anterior margin of the eye, and may extend to the mid eye. Small, larval teeth first form on the anterolateral aspect of the premaxillae at about the time the third spine of the dorsal fin is forming, these teeth disappear and are replaced by adult premaxillae teeth between 10-15 mm, except D. balteatus and G. gymnoptera which do not develop adult teeth on the premaxillae. Dentary teeth form soon after the larval premaxillae teeth (3-4 mm), these teeth are retained throughout development. Canines developed only in one C. caerulaurea larva at 18 mm and C. cuning >18 mm. Spination on the head is well developed in the least developed specimens: it is present on the preopercle in all species and Sci and Pt in some species. All head spines are smooth with the exception of serrae on the PAsp and sometimes the adjacent outer preopercular spines in three species (c. lunaris, P. chrysozona and P. tile). Head spination develops as follows: the preopercIe, ScI, Ptd and 10 spines first appear from 3 mm, Sb spines appear between 13 mm and 22 mm. Preoper- cular spines are formed in the smallest larvae and persist until about 18-22 mm. The spine at the angle of the preopercle is the largest on the head (2-15% BL), often reaching as far as the PI base and occasionally beyond. It is present in the 320 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

smallest larvae and remains conspicuous throughout the pelagic stage, The PI has as many as 17 small spines which persist to about 18 mm. The PO is armed with as many as 23 spines. All larvae smaller than 10 mm have only one POD spine and some larvae retain only a single spine to a size of 19 mm. The number of POD spines increases between 10-19 mm, but the additional spines are small and in the larger larvae perhaps better characterized as serrae. Both PI and PO spines gradually increase in number to about 18 mm, but thereafter their growth in size does not keep pace with that of the fish, and eventually a serrate border forms on the preopercle well before settlement. The single Op spine develops in larvae 3-4 mm long and is retained as part of the adult head spination. Interopercular spination forms initially as a single small spine between 3-10 mm just dorsal to and approximately parallel to the spine at the preopercular angle. Additional 10 spines may be present in some species in larvae larger than 16 mm and a serrate edge may ultimately develop. A small spine on the Sb forms from 13 mm (except no Sb spine is present in C. lunaris or P. tile); more than one Sb spine forms only in four species (D. balteatus, C. cuning, C. caerulaurea and G. gymnoptera). All the spination of the bones of the opercular series eventually disappears except the Op and that portion of the opercular series which is retained by adults in the reduced form of serrae. A small spine forms on the dorsal Pcl just dorsal to the pectoral base at 5-6 mm and persists to 25-30 mm. The Scl develops its first spine well before flexion (2 mm), and up to 3 spines form leaving a serrate edge in the largest specimens. A small Ptd spine appears at the same time as the Scl spine with two to five spines present during development; a serrate edge remains in the largest specimen (30 mm). A single Ptv spine forms between 4-6 mm and disappears between 18 mm and 23 mm. A low, initially smooth, FS ridge develops at about 3 mm. The first FS spine appears from about 4 mm and up to 10 spines develop before they reduce to serrae. The FS ridge becomes smooth again at about 20 mm, disappearing late in the pelagic period. A low, smooth F ridge dorsal and slightly anterior to the FS ridge forms at about 8 mm. A low, smooth Pe ridge forms at about 4-7 mm approximately in line with and posterior to the FS ridge, and is still present in the largest specimens. The anterior end of the maxilla develops an acute angle. Many of the spiny and smooth ridges on the head develop into the walls of the sensory canals. These include the P, FS, F, Pe and portions of the Pt. Flexion begins at about 3.7-5.0 mm and is complete at about 4.5-5.5 mm. The spines of the fins and their development are among the most distinctive charac- teristics of caesionine lutjanid larvae. The first fin elements to form are Dsp2 and P2sp. At about the time Dsp5 forms (at the beginning of flexion), the Aspl and

2 start to develop. Most of the D spines and P2 rays form in advance of the soft rays of the dorsal fin, anal fin and the primary caudal rays. However, the latter group are all present by the time notochord flexion is complete. Except for Dspl, which in some cases forms after Dsp2 or 3, the dorsal spines form from anterior to posterior. Dsp10 (Dsp1O-14 in D. balteatus) forms initially as a soft ray and transforms into a spine shortly after flexion. Asp3 forms initially as a soft ray at about 4-5 mm and transforms into a spine during or just after flexion, except in P. tile in which transformation is not complete until ca. 16 mm. P I and C procurrent rays are the last fin elements to complete formation, at about the same

time. Dsp2, P2sp and P2 ray 1 become very elongate (20-50% BL, and Dsp2 up to 60-90% BL in C. cuning, Fig. 6; see species accounts for details). Dsp2 is

initially longer than the P2 spine but they gradually become equal in length. Maximum relative spine length is reached during or shortly after flexion. The first

P2 ray is longer than the spine. Anal spines vary from being shorter to slightly READER AND LEIS: CAESIONINE LUTJANID LARVAE 321

longer than anal rays. The P2 spines in all species and the Dsp2 of G. gymnoptera, P. chrysozona and C. lunaris are trapezoidal in cross section as is Asp2 of G. gymnoptera, whereas all other fin spines are chevron-shaped in cross-section. The membrane of the spiny portion of the fins is often pigmented. Fin-spine ornamentation varies among species and developmental stage. Serrae are found on the leading edges of all the spines at some stage throughout development, but not on all trailing edges. There is no obvious internal structure in the spines. Scales form relatively late. Depending on species, incipient lateral-line scales are present from about 12-13 mm, and full complements of scales appear from 16-17 mm during the pelagic period. Pigment.-Larvae are initially lightly pigmented but acquire more pigment as transformation approaches. A pigment cap is present dorsally on the gas bladder and gut. Pigment does not appear dorsally on the brain until approximately 5 mm when the first melanophores form on the midbrain. Initially one or more melanophores are present along the ventral midline of the tail, but the subsequent fate of these melanophores varies among taxa. A series of melanophores originates on the dorsal midline of the caudal peduncle after flexion and spreads onto the D base in larvae larger than 10 mm. Pigment appears on the urostyle soon after flexion (during flexion in P. tile). Often pigment is present on the D and P2 membrane in association with the fin spines and/or in the spine chevron. Mela- nophores may be present at the cleithral symphysis only over limited size ranges. Other pigment varies among taxa.

Species Accounts of Development Genus Caesio Caesio caerulaurea (Lacepecte, 1801)--42 specimens, 3.75-28.13 mm Figures 2-7, Table 3

Material Examined.-East China Sea, AMS 1.31832-001, 1.31979-002, 1.31979-003, 1.31998-001, NSMT-PL 32-35: Bismarck Sea, AMS I.19713-028, I.19726-030 to -033: Solomon Sea, AMS I.19741- 035: Coral Sea, AMS 1.23133-063, 1.23551-009, 1.24211-010, 1.24524-002, 1.24572-002. 1.24617-003, 1.26617-005, 1.30881-025 to -028, MCZ 110028, MCZ 110029. Diagnosis-MORPHOLOGY. Fin spines long (MRL of Dsp2 47% BL shortly after flexion, Fig. 2); fine to moderate serrae on single leading edges of fin spines; PI border with 9 spines to 18 mm; one 10 spine to 11 mm, 3-11 at 18-22 mm; Sb 1-5 at 20-28 mm; PAsp smooth and long (MRL 14% BL, Fig. 5), reaching beyond pectoral base to 10 mm; D X,14-16, A m,12-13. PIGMENT.Flexing larvae with two melanophores on the ventral midline of the tail, on the posterior anal anlage and caudal peduncle (Fig. 2A); no pigment dorsally on trunk or tail until 6.0 mm; chevron pigment on Dsp2 in least developed larva and on P2sp at 4.0 mm; anterior CS pigment at 3.7 mm, first absent at 4.8 mm, consistently absent at 6.0 mm; no preanal median finfold pigment. Description.-MoRPHOLOGY. Body moderately deep (27-39% BL); head moderate to large (31-44% BL); snout (8-14% BL) shorter than maxilla (9-17% BL), which falls short of anterior edge of pupil. Larval teeth on premaxillae tip and dentary in least developed larva; two canines at the tip of the premaxillae at 18 mm. Head spines smooth. PAsp reaching beyond PI base in specimens to 10 mm (MRL 14% BL, Fig. 5); from 18 mm (the next largest specimen) PAsp reaching to or just short of PI base. Preopercu1ar spines moderate to large, the least developed specimen (3.75 mm) with no spines on PUI (PUI 0); PLI 1-2; PUO 1; PLO 2-3; number of spines increasing to maximum of PUI 4; PLI 6; PUO 1; 322 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

Figure 2. Larvae of Caesio caerulaurea from the East China Sea. Scale bars = 1 rom, A. 3,8 mm (NSMT-PL 33); B, 5.3 rom (AMS 1.31832-001),

PLO 7 at 10 mm, at 18 mm to the largest larva PI is smooth and PO serrate. 10 1 at 3.75 mm; between 3 and 11 at 18-28 mm. Sb between 1 and 5 at 20-28 mm. ScI 1 in the least developed larva; 2 at 3.8 mm; 3 at 6.8 mm; reduced to 1 at 19 mm and a serrate edge in the largest specimen. Ptd 1 at 3.8 mm; 2 at 7.3 mm; 1-4 in larvae 19-28 mm. One Ptv in 6 of 14 larvae 3.8-6.3 mm, one present in all larvae thereafter, disappearing at 23 mm. FS smooth in preflexion larvae; READER AND LEIS: CAESIONINE LUTJANID LARVAE 323

c Figure 3. Larvae of Caesio caerulaurea. Scale bars = I mm. A. 8.0 mm from the Bismarck Sea. (AMS 1.19726-032). B. 10.7 mm from the western Coral Sea. (MCZ 110028). C. 19.1 mm settled larva from the Great Barrier Reef. near Lizard Island (AMS 1.30881-025). Specimen is fully scaled but scales shown only on lateral line. 324 BULLETIN OF MARINE SCIENCE, VOL. 59. NO.2, 1996

Figure 4. Larva of Caesio caerulaurea. Scale bar = I mm. 28.1 mm settled larva from Great Barrier Reef, near Lizard Island (AMS 1.24617-002). Specimen is fully scaled but scales shown only on lateral line.

1 at 4.7 mm; 1-8 weak spines in larvae 4.8-18 mm; smooth at 19 mm. Weak F at 8-9 mm to the largest specimen. Weak Pe at 3.8 mm to the largest specimen. Notochord flexion occurs at 3.8-4.7 mm. In the least developed specimen Dspl-4, Aspl, P2sp and four P2 rays present; at 6.0 mm all fin elements present except PI and procurrent C rays; PI and procurrent C rays complete and all fin spines fully ossified at 7-8 mm. Leading edges of Dspl-4 with serrae, remaining leading edges smooth, all trailing edges serrate. At 18 mm, serrae begining to disappear; all D spines smooth at 25 mm. Aspl-2 with serrate leading and trailing

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Figure 5. Caesio spp. Length of preopercular angle spine (as a percent of body length) in relation to body length. READER AND LEIS: CAESIONINE LUTJANID LARVAE 325

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Figure 6. Caesio spp. Length of dorsal-fin spine 2 (as a percent of body length) in relation to body length.

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45 0 40 r!§!Jo~ @ 0 00 ~O o"g o 0 35 • ••• •• 30 • • 0 .ll* + ~ + • • ~ • + !g 25 0+'* 0 Q. (/J 20 0 C\I cF a. 15 o 10 o o 5+-..,-..,-..,-..,-..,-..,-..,-..,-..,-..,-..,-..,-,---,---,---,---,---,---,---,----r-i 2 4 6 8 10 1'2 14 16 18 20 22 24 Body Length (mm)

• C. caerulaurea + C. lunaris o C. cuning

Figure 7. Caesio spp. Length of pelvic spine (as a percent of body length) in relation to body length. 326 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2. 1996

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Remarks.-The series has a large gap between 10 and 18 mm, a size range over which a considerable number of pigment and morphological changes take place. Very small larvae are also lacking. Larvae of the three Caesio spp described here differ in a number of characters, and while it is not possible to confidently predict the distribution of these characters among the other 5 Caesio spp, we think that it is likely they will resemble C. caeru/aurea more than the other two species. Our reasoning for this statement is that neither the distinctive fin-spine morphology of C. cuning. nor the serrate PAsp of C. /unaris have been seen in any Caesio or Pterocaesio larvae we were unable to identify to species. READER AND LEIS: CAESIONINE LUTJANID LARVAE 329

Caesio cuning (Bloch, 1791) 50 specimens, 2.7-23.9 mm Figures 5-10, Table 4

Material Examined.-Gulf of Thailand, USNM 330431: western Coral Sea AMS 1.20990-063, 1.23110-008,1.23113-022,1.23130-018,1.23131-023, 1.23131-024, 1.23131-025, 1.23131-026, 1.23133- 028, 1.23133-029, 1.23133-030, 1.23133-031, 1.23133-032, 1.23133-033, 1.23133-062, 1.23513-004, 1.23579-011. 1.23592-005, 1.23742-003, 1.23742-004, 1.23760-004, 1.23760-005, 1.23760-006, 1.23760- 007, 1.24196-008, 1.24208-014, 1.24208-015, 1.24208-016, 1.24209-012, 1.24209-013, 1.24209-033, 1.24209-034, 1.24748-002, 1.24748-004, 1.24784-003, 1.26347-004, 1.26350-007, 1.26358-002, 1.34163-001, 1.35443-001 to 003, 1.35444-001, 1.35445-001: Bougainville Is., Papua New Guinea, CSIRO, L 58, L 65. Diagnosis.-MORPHOLOGY. Fin spines very long (MRL of Dsp2 94% BL shortly after flexion, Figs. 6, 8) with serrae on the single leading edge and fine comb- like serrations on trailing edges; up to 11 spines on PI border; 1 spine on 10 from 3.6 to 17 mm, 1 to 8 present from 17 mm; 1 spine on Sb at 12 mm; PAsp with auxiliary spines at 21 mm, moderately long, reaches to PI base before 12 mm BL (MRL 12% BL, Fig. 5); D X,14-16, A III,10-12. PIGMENT.Initially three melanophores on the ventral tail, from just posterior to the anal anlage to the fin fold near the caudal anlage (Fig. 8A); no pigment dorsally on trunk or tail until after flexion (6.7 mm); Dsp2 and P2sp with chevron pigment inconsistently from 3.] mm, expanding to Dsp3 and associated membrane from 20 mm, and expanding over adjacent P2 rays and membrane from 6.7 mm; melanophore present on anterior edge of CS from 2.8 mm to 6.6 mm; pigment on the preanal median finfold in a few preflexion specimens. Description.-MoRPHOLOGY. Body moderately deep (25-38% BL). Head moderate to large (23-42% BL); snout (8-]5% BL) equal to or slightly shorter than maxilla (8-17% BL); maxilla reaching to or just beyond anterior edge of pupiL Larval teeth present on premaxillae at 3.4 mm and dentary at 3.4 mm (Fig. 8A); canines present on tip of premaxillae at ] 8 mm. Head spines smooth except for auxiliary spines on PAsp in one 2] mm larva (Fig. lOB). PAsp (MRL ] 1% BL) reaching base of PI to 12 mm, from 16 mm (the next specimen in the series) PAsp falls well short of PI base. Preopercular spines moderate to large, the least developed specimen (2.7 mm) with PUll; PLI 2-3; PUO 1; PLO 2-3. Number of spines increasing to a maximum of PUI 5 at 9.4 mm; PLI 7 at 7.6 mm; pu~ ]4 at 18 mm; PLO 8 at 12 mm. PUI and PLI spines absent at 18 mm; PUO and PLO spines becoming much reduced, ultimately remaining as serrae. 10 ] at 3.6-- 16 mm; between 4 and 10 spines may be present from ]7-24 mm. Sb ] at 12 mm; from 0 to 3 may be present in larvae to 24 mm. ScI 1 at 2.7 mm; 2 at 4.0 mm, 2-4 present thereafter. Ptd 1 at 3.1 mm; 2 at 5.3 mm, 1-6 from 11 mm to 24 mm. Ptv 1 at 5.0 mm; beginning to disappear at 17 mm, absent at ]8 mm. FS smooth at 2-3 mm; 1 at 3.8 mm; 8-10 weak spines between 8.5-11.7 mm; thereafter reduced to small serrae; smooth at 21 mm. Weak F at 8-9 mm to the largest specimen. Weak Pe at 5.2 mm, becoming stronger with size to the largest specimen. Notochord flexion occuring at 4.7-5.4 mm. In the least developed specimen

Dsp2, P2sp and one P2 ray present; at 6.1 mm all fin elements except PI and procurrent C rays present; a full complement of PI and procurrent C rays at 7.8 mm. Dsp2 10-92% BL; P2sp (10-50% BL) usually half length of Dsp2; MRL of Dsp2 and P2 at 6-7 mm (Figs. 6, 7). In preflexion larvae leading and trailing edge of Dsp2 serrate but only trailing edge of Dsp 1 with serrae. Leading edges of Dspl-3 with serrae soon after flexion, leading edge of Dsp3 smooth again at 7.0 mm, other Dsp leading edges smooth throughout. Trailing edges of all Dsp with fine comb-like serrae at 9.6 mm; serrae beginning to reduce in size at about 330 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

Figure 8. Larvae of Caesio cuning from the Great Barrier Reef, near Lizard Is]and. Scale bars = 1 mm. A. 3.4 mm (AMS 1.23133-034). B. 4.2 mrn (AMS 1.23] 10-008). C. 5.4 mm (AMS 1.23592-005).

10 mm; smooth at 23 mm. Asp1-2 with serrae on leading and comb-like serrae on trailing edges soon after flexion; Asp3 leading edge smooth to 9.8 mm thereafter serrate distally, trailing edge serrate at 1] mm; serrae beginning to disappear at 16 mm, absent by 19 mm. P2SP with serrate leading and trailing edges until 20 mm; thereafter smooth. Comb-like serrae on fin spines confluent with comb-like READER AND LEIS: CAESIONINE LUTIANID LARVAE 331

Figure 9. Larvae of Caesio cuning. Scale bars = ] mm. A. 7.6 mm from the Great Barrier Reef, near Lizard Island (AMS 1.24208-015). B. 10.4 mm from the Gulf of Thailand (USNM 33043]). internal structure of the spines (Figs. 1, 8-10); internal structure absent in the largest specimen, but only well after the disappearance of the external serrae. At 11 mm, no scales, by 17 mm completely scaled, but due to the gap in the series between the two, size at onset of scale formation unknown. PIGMENT. In the least developed specimen three ventral melanophores on the tail, posteriormost on fin fold near caudal anlage (Fig. 8A); during flexion, one at posterior A anlage, one at mid caudal peduncle, and one at C anlage. At about 9 mm ventral pigment on tail increasing anteriorly along peduncle and A-ray bases; at 20 mm pigment extending entire length of A base and peduncle. Me- lanophore at base of last D ray first present at 6.7 mm, consistently present at 8.5 mm along with a second melanophore at mid peduncle; dorsal pigment following the same configuration as ventral pigment, from 17 mm along D base anteriorly and posteriorly to mid peduncle. At flexion, a melanophore on lower mid C base; 332 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

Figure 10, Larvae of Caesio cuning. Scale bars = 1 mm. Specimens are fully scaled but scales shown only on lateral line. A. 17.7 mm from the western Coral Sea (AMS 1.24748-003). B, 20.6 mm larva from Bougainville Is. (CSIRO L58). Fine serrae are still present on Dsp2 and 3 but not illustrated. Note serrations on PAsp. Hind brain pigmented though hidden by myomeres. from 9.6 mm pigment increasing over lower C ray bases; at 17 mm pigment covering all lower principal C ray bases; and at 19 mm pigment present on upper C ray bases. Internal melanophore over urostyle at 6.9 mm; melanophores increasing in number from 9.4 mm; at 17 mm a cluster of up to 10 melanophores present over the urostyle. Melanophore anterior to CS in least developed larva; first absent at 6.7 mm; last present at 8.3 mm. A melanophore on preanal medial finfold in four of fourteen preflexion larvae. Ten to 16 dorsal melanophores on the midbrain at 6.1 mm; complete cover at 17 mm. Two internal melanophores at the base of the hindbrain at 6.6 mm; at 19 mm hindbrain completely covered though hidden by myomeres. A melanophore on forebrain dorsally first present at 8 mm; consistently present from 11 mm; forebrain completely covered at 17 mm. Four melanophores on premaxillae at 8.3 mm; inconsistently present to 11 mm; after 17 rom pigment scattered dorsally on snout, premaxilla, dentary and under chin. One internal melanophore at Op spine base first present at 6.7 mm, consistently present from 8.5 rom. At 20 mm scattered lateral pigment on trunk and tail and over outer two thirds of the caudal fin (Fig. lOB). A series of me- READER AND LEIS: CAESIONINE LUTJANID LARVAE 333 lanophores in chevron and adjacent membrane of Dsp2 and in distal portion of chevron and adjacent membrane and ray tip of P2sp first present at 3.1 mm, consistently present from 4.8 mm. At 23 mm melanophores over membrane of

Dspl-7. At 17 mm P2 heavily pigmented especially along longest ray; pigment disappearing by 23 mm. Remarks.-The series of C. cuning is the most complete of the three Caesio spp. and has the greatest size range, but still there is a gap between II and 17 mm. Caesio cuning has by far the longest Dsp2 of any caesionine, and very distinctive comb-like ornamentation on its fin spines. However, this ornamentation is very similar to that found in Pterocaesio tile. Larvae of the two species may be identified by the differences in spine length, meristics and pigment (Tables I, 2). Caesio cuning larvae have one of the more "coastal" distributions found amongst caesionines, with nearly all larvae being found over the continental shelf, and most over the mid to inner portion of the shelf (I.M.L., unpubl.). Pterocaesio tile larvae, in contrast, have perhaps the most oceanic distribution among caesionines (I.M.Leis., unpubl.). This correlates well with adult distribution: C. cuning is not found on oceanic reefs, and is restricted to continental shelves west of the An- desite Line (Springer, 1982), whereas P. tile is widely distributed on oceanic reefs in the Indian and Pacific Oceans including the Pacific Plate (Carpenter, 1987, 1988).

Caesio lunaris (Cuvier, 1830)-16 specimens, 4.5-9.1 mm Figures 5-7, 11, 12, Table 5

Material Examined.-East China Sea, AMS 1.31540-031, 1.31633-001, NSMT-PL 36: Philippine Sea, AMS 1.31527-002: western Coral Sea, AMS 1.30536-001, 1.30536-002, MCZ 110030, MCZ 110031; eastern Indian Ocean over northwest shelf of Australia, AMS 1.35037-001; Andaman Sea, AMS 1.32034-002; Banda Sea, ZMA 121.163-164. Diagnosis.-MORPHOLOGY. Fin spines long (MRL of Dsp2 50% BL shortly after flexion, Fig. lIB) with small to moderate serrae on leading and trailing edges; Dsp2 with two leading edges; up to 17 spines on PI border; I spine on 10 and 0 on Sb; PAsp long, reaching to just beyond PI base (MRL 15% BL, Fig. 5); PAsp serrate; D X, 13-15, A III,IO-11. PIGMENT.Two melanophores on the tail ventrally in postflexion larvae, one at the base of last two A rays and one at mid-peduncle; a melanophore at the base of the last two D rays (Fig. lIA); chevron pigment on Dsp2 from 4.9 mm and Dsp3 from 5.5 mm, both inconsistent thereafter; chevron pigment on distal half of P2 and tip of first ray at 4.9 mm, inconsistent thereafter; pigment anterior to the CS in all specimens smaller than 5.0 mm, inconsistent to 6.8 mm thereafter absent except for one 8.3 mm specimen. No preanal median finfold pigment. Description.-MoRPHOLOGY. Body moderate to deep (32-41 % BL); head moderate to large (36-46% BL); snout (10-16% BL) shorter than or equal to maxilla (10-16% BL); maxilla initially reaching to or just short of anterior edge of pupil. Larval teeth on tip of premaxillae and adult teeth on dentary at 4.5 mm. PAsp reaching beyond PI base (MRL 15% BL); all head spines smooth except PAsp which has small serrae in all larvae. Preopercular spines large, the least developed larva with PUll; PLI 3-4; PUO 1; PLO 4 (Fig. IIA). Number of spines increasing with size to a maximum of PUI 4 at 8.1 mm; PLI 6; PLO 6 at 9.1 mm (Fig. 12B). 10 1; Sb 0; Scl 2 at 4-5 mm; a third appearing first at 5.4 mm, consistently from 6.1 mm. Ptd 1 in the least developed specimen; 1-2 at 5-9 mm. Ptv 1 at 4.5 mm. FS 1-4 between 4.5 mm and 7.5 mm; 5 (weak) at 8.1 mm; 6 (weak) at 334 BULLETIN OF MARINE SCIENCE. VOL. 59. NO.2. 1996

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Figure 11. Larvae of Caesio lunaris. Note serrate PAsp. Scale bars = 1 nun. A. 4.5 nun from East China Sea (NSMT-PL 36). B. 5.4 nun from the Andaman Sea (AMS 1.32034-002).

9.1 mm. F absent. Pe weak at 5.4 mm. Notochord flexion complete in smallest specimen (4.5 mm). At 5.4 mm all fin elements present except PI and procurrent C rays, ossification of all fin spines and procurrent C rays complete but PI incomplete in largest larva. Dsp2 and P2sp short; Dsp2 32-50% BL; P2sp 25-35% BL. The first P2 ray slightly longer than the spine. Dsp2 with two leading edges in all larvae. Leading edge(s) of Dspl-3 with serrae from the least developed larva, and Dsp4 in 4 of 11 larvae between 5.4-9.0 mm; remaining Dsp leading edges smooth. Trailing edges of Dspl-4 serrate in the least developed larva; all trailing Dsp edges serrate at 9.1 mm. Aspl and 2 with serrate trailing edges in the least developed larva, and serrate leading edges at 5.4 mm; Asp3 smooth to 8.0 mm thereafter serrate on all three edges. A spines with single leading edges. Scales absent in largest specimen. READER AND LEIS: CAESIONINE LUTJANID LARVAE 337

Figure 12. Larvae of Caesio /unaris. Note serrate PAsp. Scale bars = I mm. A. 7.5 mm from the western Coral Sea (MCZ 110030). B. 9.1 mm from the Banda Sea (ZMA 121.164).

PIGMENT.The least developed larva with two ventral melanophores on tail, one at base of last two anal rays, and one centrally on peduncle; two melanophores at the posterior two D ray bases (Fig. lIA). Melanophore laterally on lower portion of C peduncle in two larvae of 6-7 mm. Melanophore at base of first to third lower C rays from 4.5 mm. Internal pigment patch on urostyle in all larvae, becoming larger with size. Pigment consistently present anterior to CS between 4.5-5.0, inconsistently present to 8.3 mm, thereafter absent. Internal pigment on the hindbrain base at 5.2 mm; 2-12 melanophores (both internally and externally) over top and posterior surface of midbrain at about 5 mm, increasing in number to almost cover midbrain at 9.1 mm. Forebrain with one to two melanophores dorsally from 8 mm. One to three melanophores each on premaxillae and lower jaw tip from 6.8 mm. Internal melanophore at base of Op spine from 5.4 mm.

Dsp2 (spine and adjacent membrane) and P2sp (spine, adjacent membrane and 338 BULLETIN OF MARINE SCIENCE. YOLo 59. NO.2. 1996

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first ray) chevron pigment first present at 4.9 mm, thereafter on P2sp in 7 of 13 larvae and on Dsp2 in 9 of 13 larvae. Remarks.- The rather limited series of C. lunaris (no preflexion larvae, and none larger than 9.1 mm) limits the description we can supply. The most distinctive feature of this species is the serrate PAsp in all specimens. The F ridge is probably present in larvae larger than the largest in our series.

Genus Dipterygonotus (monotypic) Dipterygonotus balteatus (Valenciennes, 1830)-87 specimens, 2.4-21.8 mm Figs. 13-16, 17-19, Table 6

Material Examined.--Celebes Sea, ZMUC DANA St. 3738-II1: Bismarck Sea, AMS 1.19694-015: western Coral Sea, AMS 1.20948-036, 1.22548-006, 1.23053-072, 1.23201-011, 1.23742-005, 1.23746-001, 1.23754-006, 1.23755-001, 1.23764-006, 1.24511-003: western Tasman Sea, AMS 1.26956-003, 1.29907-001,1.33842-001: eastern Indian Ocean over northwest shelf of Australia, AMS 1.26415-001, 1.26415-002,1.26415-003,1.26416-001,1.26417-001, 1.26417-002, 1.26417-003, 1.26417-005, 1.26418- 001, 1.26418-002, 1.26419-001, 1.26420-001, 1.26421-001, 1.26422-001, 1.26423-001, 1.26424-001, 1.31950-002,1.31956-003,1.33814-002,1.33830-002, 1.33838-002, 1.33839-002, 1.33843-001, 1.33844- 001, 1.33845-001, 1.33846-001, 1.33847-001, 1.33848-001, 1.33849-001, 1.33850-001, 1.33851-001, 1.33852-001, 1.33853-001, 1.33854-001, 1.33855-001: eastern Indian Ocean west of Sumatra, AMS 1.32513-003: Andaman Sea, AMS 1.32398-004. Diagnosis.-MORPHOLOGY. Fin spines moderately long (MRL of Dsp2 36% BL shortly after flexion, Fig. 18) with moderately course serrae on leading and trailing edges; single leading edges on all spines except P2; many spines on PI border (up to 17); up to 9 10 and 5 Sb spines; PAsp moderately long and smooth, reaching to or beyond pectoral base in all but the largest larva (MRL 12% BL shortly after flexion, Fig. 17); D XII-XV,8-11, A III,9-11. PIGMENT.Initially four melanophores on ventral midline of tail, from just pos-· terior to anus to caudal anlage (Fig. 13A); three remaining in postflexion larvae:: no pigment dorsally on trunk or tail until just after flexion; D spines and P2sp with pigment; one melanophore anterior to CS from 2.8 to 6.6 mm; no melano·· phore on the preanal median finfold. Description.-MoRPHOLOGY. Body depth moderate, initially 35% BL, becoming more slender with size to 21% BL at 21 mm. Head moderate to large (26-42% BL); snout (8-15% BL) shorter than maxilla which may reach to anterior edge of the pupil. Teeth present on premaxiIlae from 2.8 mm and tip of dentary soon after. From about 11 mm premaxiIlary teeth decreasing in number, disappearing by 15 mm. Dentary teeth still present in the largest specimens. No canine teeth present. Head spines large and smooth. PAsp (MRL 12% BL) reaching to beyond PI base in least developed larva (2.4 mm), to PI base in all other larvae except largest where it falls short of PI base. Preopercular spines large, forming early, the least developed larva with PUll; PLI 3; PUO 1; PLO 2. Number of preopercular spines increasing with size, to a maximum of: PUI 10 at 13 mm; PLI 9 at 11-14 mm; PUO 8 at 17 mm; PLO 12 at 16 mm. Thereafter, number of preopercular spines decreasing; all spines but PLI present in the largest specimen. 10 1 at 4.3 mm (flexion); 2 at 12 mm; up to 6 at 15 mm; largest larva with 9 10 spines. Sb 1 at 16 mm, up to 5 at 21 mm. ScI 1-2 at 3 mm (preflexion); 3rd at 5.7 mm; thereafter a 4th may form. Ptd 1 at 3.2 mm to flexion; up to 4 at 11 mm and 5 at ]5 mm. Ptv ] from 4.1 mm to 18 mm, absent thereafter. FS smooth at 3.2 mm; 1st serrae at 4.1 mm; ] 1 at 13.3; 8-9 at 18 mm; no serrae at 20 mm but some irregular bumps still present. F present from 8.0 mm. Pe bump at 5.8 mm; forms low, weak ridge soon after; strongest at 20 mm. Notochord flexion occUJr- 340 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

Figure 13. Larvae of Dipterygonotus balteatus from the Great Barrier Reef, near Lizard Island. Scale bars = 1 mrn. A. 2.4 mrn CAMS 1.23053-072). B. 2.8 mrn CAMS 1.23053-072). C. 3.2 mrn CAMS 1.23764-006). READER AND LEIS: CAESIONINE LUTJANID LARVAE 341

Figure 14. Larvae of Dipterygonotus balteatus. Scale bars = 1 mm. A. 4.1 mm from the Great Barrier Reef, near Lizard Island (AMS 1.23201-011). B. 4.8 mm from the eastern Indian Ocean, over the North West Continental Shelf of Australia (AMS 1.26417-005).

ring at 4.2-5.1 mm. All D, A and P2 fin elements present just after flexion (5.8 mm). Posteriormost A and four D spines initially forming as rays but ossifying to spines just after flexion. All PI rays present at 7-8 mm. Dsp2 present in least developed specimen, shorter than P2 spine until 3.2 mm, thereafter both spines approximately equal in length. P2sp and Dsp2 (16-30% BL) reaching MRL at 4-

9 mm (Figs. 18, 19). All fin spines except P2 with one leading edge. Fin spines with moderately course serrae; these weakening from 13 mm and absent in largest specimen (22 mm). Incipient scales on the cheek and anteriorly on the lateral line at about 13 mm; full complement of incipient lateral-line scales at 14 mm; scales complete at 16 mm. PIGMENT.Four melanophores evenly spaced on ventral midline of tail until flexion, the first anteriorly on A anlage and last on ventral notochord tip at caudal anlage (Fig. 13A). Following flexion one melanophore present at base of A ray 2, one just posterior to ultimate A ray, one midway along peduncle and one at 342 BULLETIN OF MARINE SCIENCE. VOL. 59. NO.2, 1996

Figure 15. Larvae of Dipterygonotus balteatus. Scale bars = 1 mIll. A. 6.5 mIll from the Andaman Sea (AMS 1.32398-004), g, 7.4 mIll from the western Coral Sea (AMS 1.24511-003). base of lower C rays (Fig. 14B). From 7.0 mm, melanophores on ventral midline of tail increasing in number; largest specimen with seven melanophores along A fin base and a continuous line of melanophores on peduncle from A to C. Pigment dorsally on tail initially at 5.7 mm, extending to mid peduncle from D base and along the entire peduncle by 10.7 mm; at 5.7 mm three melanophores at base of Drays 1-3; at 6.5 mm additional melanophores present at base of Dsp 12-14; largest larva with melanophores from base of Dsp3 to end of D fin. At 9-10 mm melanophores increase at base of lower C rays, spreading to upper C ray bases by about 11-13 mm. One melanophore internally on urostyle dorsally (concave surface) just after flexion (5.0 mm), increasing to a small cluster at 11 mm remaining thereafter. Single melanophore anterior to CS on ventral midline, con- READER AND LEIS: CAESIONINE LUTJANID LARVAE 343

c Figure 16. Larvae of Dipterygonotus balteatus from the eastern Indian Ocean over the North West Continental Shelf of Australia. Scale bars = I mm. A. 11.2 mm (AMS 1.33844-(01). B. 13.3 mrn (AMS 1.33838-002). C. 20.5 mm (AMS 1.26416-(01). Specimen is fully scaled, but scales are shown only along lateral line.

sistently present to 4.6 mm, inconsistently present to 7.5 mm, thereafter absent. A melanophore posterior to CS in larvae smaller than 3 mm (Fig. 13A, B). No pigment on the preanal median finfold. An internal, posterior melanophore on hindbrain ventrally at 2.8 mm (Fig. 13B); after flexion internal pigment on hindbrain increasing, extending to junction of mid- and hindbrain and ventrally under hindbrain. External pigment dorsally on midbrain at 5.0 mm and forebrain at 8.1 mm; at 15 mm, dense cover of melanophores present dorsally over both fore- and midbrain externally and hindbrain internally. At 6.5 mm one to two dorsal me- 344 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

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'0 Q) :l ~~~~~~~~~~8~~~~~~~~~~~~8 .5 oooooooo~oo~~~oooooOOOO~~NNNN E o ------u ~~~NOOOOO~OOOO~OOOO~NOON~OO~OO~ N~~~OOOON~~~~OOOON~~O~-NNOO~~ ------NN~~~~~~~~~~~~~~~~OOOO~~~~O~ READER AND LEIS: CAESIONINE LUTJANID LARVAE 347 lanophores on snout (Fig. 15A), number increasing with size. At 7.7 mm mela·· nophores present on premaxillae and dentary symphysis; at 15 mm melanophores laterally and ventrally on dentary. An expanded, external melanophore at base of opercular spine during flexion (4.8 mm); a second may appear at about 8 mm; thereafter, 1-8 melanophores in this region. Four of six larvae at 19-22 mm with 6-7 small melanophores around the posterior border of orbit. One to several melanophores present in the chevron of P2 and Dsp2 spines at 2-3 mm (except in the two least developed larvae); Dsp2 melanophores associated with adjacent membrane; P2sp melanophores associated with adjacent membrane and ray. P2 pigment reducing with size, absent at 20 mm. At 15-21 mm, melanophores spreading to Dsp3-6. From 19 mm, small, external melanophores scattered dorso- laterally above the lateral line (Fig. 16C). Remarks.-Preflexion P. chrysozona larvae are very similar to D. balteatus. How- ever, a single melanophore is present at the preanal median finfold in small preflexion P. chrysozona: this pigment is absent in D. balteatus. In the larger preflexion and all postflexion larvae Dsp2 has a single anterior (or leading) edge in D. balteatus whereas Dsp2 in P. chrysozona has a double anterior edge. Addi- tional material of D. balteatus from the western Indian Ocean, Andaman Sea and Gulf of Thailand, not included in the above description, did not notably differ from the specimens upon which our description is based. D. balteatus is among the most abundant percoid larvae encountered over open continental shelf habitats in the eastern Indian Ocean (Andaman Sea and Austra- lia), particularly in winter. Therefore we have been able to assemble a particularly complete series. The larvae seem less abundant in equivalent habitats in the Pa- cific, but this could be an artifact of sampling effort. The great abundance of these larvae indicates a great abundance of adults, but given the small size of aduhs (about 140 mm), it is unlikely they would be of great commercial importance. They are, however, used as baitfish in some areas. Collections of D. balteatus larvae examined during this study considerably extend the known range of the species. We have seen larvae from the western Tasman Sea (35°S, off Sydney, Australia) and the western Indian Ocean (34°5, off Port Elizabeth, South Africa) where the southernmost records were previously 15°S and 50S, respectively (Carpenter, 1987, 1988). Both areas are subject to strong, warm, poleward currents, so these temperate-subtropical occurrences of larvae may simply represent expatriation, but it is also possible that the relatively inconspicuous adults occur outside the tropics as well.

Genus Gymnocaesio (monotypic) Gymnocaesio gymnoptera (Bleeker, 1876)-57 specimens, 7.0-30.1 mm Figures 17-19,20,21, Table 7

Material Examined.-Eastern Indian Ocean off west coast of Sumatra, AMS 1.34606-001, 1.34607- 001 to -005, 1.34608-001, 1.34608-002, 1.34609-001, 1.34727-001; ZMUC DANA 3861-I,II,V,VI, 3867- I,II,III,IV, 3868-1, 3869-II,V,VIII; Jacquinot Bay, New Britain, CSIRO L 77. Diagnosis.-MoRPHOLOGY. Fin spines moderately long (MRL of Dsp2 35% BL, Fig. 18), with medium to fine serrae on leading and trailing edges; Dsp2 with two leading edges from 7.0 mm (the least developed specimen), Asp2 with two leading edges from 11 mm; PI border with up to 12 spines; up to four spines on the 10; one Sb spine at 21 mm, five at 30 mm; PAsp without serrae, moderately long, reaching PI base to 11 mm, thereafter falling short (MRL 13% BL, Fig. 17); D X-XI,14-16, A m,ll-13. 348 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

13 ++ 12

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Figure 17. Dipterygonotus balteatus and Gymnocaesio gymnoptera, Length of preopercular angle spine (as a percent of body length) in relation to body length.

PIGMENT.Two melanophores each on ventral and dorsal midlines of the tail in the least developed larvae, one each at base of the last D and A fin rays and one each on the dorsal and ventral margins of peduncle (Fig. 20A); Dsp2 chevron pigment present in the least developed larva; pigment on chevron and associated membrane of Dspl-IO in largest specimen; P2 pigment associated with ray I and adjacent membrane to 20 mm, thereafter absent.

Description.-MoRPHOLOGY. Body moderately deep to elongate, initially 25% BL becoming more slender with size to 21% BL at 30 mm. Head moderately large, (29-37% BL); snout equal to or longer than maxilla (9-11 % BL); maxilla reaching the orbit but never extending beyond anterior edge of eye. Teeth on premaxillae and tip of dentary in least developed specimen; premaxillary teeth decreasing in number at about 11 mm, disappearing by 15 mm; dentary teeth still present in the largest specimen; no canines. Head spines smooth; PAsp reaching PI base to 11 mm (MRL 9% BL) thereafter falling short of PI base. Preopercular spines moderately long, the least developed specimen (7.0 mm) with pur 2-3; PLI 4; PUO 1; PLO 5 (Fig. 20A). Number of preopercular spines increasing with size, to a maximum of pur 5, PLI 6 and PLO 7 at 14 mm; PUO remains at 1; preopercular spines becoming weak and much reduced from 16 mm; at 30 mm preopercular edges serrate (Fig. 21B). ro 1 at 7.0 mm; 4 at 21 mm; 3 at 30 mm. Sb 1 at 21 mm; 5 at 30 mm. Scl 2 at 7.0 mm; 3 at 7.7 mm; number of spines decreasing from 19 mm; 0 at 30 mm. Ptd 1 at 7.0; 2 at 7.7 mm; 3-4 at 18 mm. Scl and Ptd bones serrate at 30 mm. Ptv 1 at 7.0 mm; 0 at 20 mm. FS 5 serrae at 7.0 mm; 7 at 10 mm; 9 at 12 mm; 10 weak serrae at 13 mm; smooth at 23 mm. Weak F and Pe ridges at 10 mm to the largest specimen. All fins except PI READER AND LEIS: CAESIONINE LUTJANID LARVAE 349

35 • + .c. 0) • -t: .3 30 .<1II.f + ;>. •• ",..+ "'0 + 0 • • •••• m 25 • .+~...• '#. • •• gj • .-+,.i¥ +.;...• (\J 20 + a. +. (/) ~+ 0 • ;.~.+ •• + 15 • + 10 -f.=-r---r---,r--.--.--.--.----.----.---.--.---.----.------r----.-r--,----r---,-.--.---r-i 2 4 6 8 10 12 14 16 1'8 20 22 24 Body Length (mm)

• D. balteatus + G. gymnoptera I

Figure 18. Dipterygonotus balteatus and Gymnocaesio gymnoptera. Length of dorsal-fin spine 2 (as a percent of body length) in relation to body length. and C procurrent rays fully formed in least developed larvae. PI and procurrent caudal rays complete at 9-11 mm. Fin spines of moderate length (Dsp2=P2SP); both P2 and Dsp2 reach MRL of 30-35% BL at 7-8 mm; 12% BL at 30 mm. P2 ray I equal to or slightly longer than P2 spine. Two leading edges on Dsp2 at 7- 23 mm and on Asp2 at 11-23 mm; Dspl-6 with serrate leading edge, leading edge of remaining spines smooth; all D spines with serrate trailing edges at ]l2 mm; at 30 mm all spines smooth except for weak serrae at base of trailing edge of P2sp. Incipient scales anterodorsally on the trunk in larvae as small as 13 mm; complete scalation at 30 mm. PIGMENT. In the least developed specimen (7.0 mm) a melanophore at posterior base of D and A fins; one each anteriorly on dorsal edge and centrally on ventral edge of peduncle (Fig. 20A). At 10 mm small melanophores along base of D and A rays. In 4 of 17 specimens (16-20 mm) melanophores at base of Dsp3-1O. At 9.6 mm and larger, peduncle melanophores increasing on ventral edge and forming paired line along entire dorsal edge. Initially a melanophore at second lower C ray base (Fig. 20A); at 10-12 mm one to two melanophores on bases of upper rays; from 14 mm melanophores present over middle C ray bases. An internal melanophore on dorsal side (concave) of urostyle in least developed specimen (Fig. 20A). In the largest specimen (30 mm) urostyle pigment merging with a band of mid-lateral pigment spots (Fig. 21B). No CS pigment. Six internal melanophores on midbrain dorsally at 7.0 mm, the number increasing from 10 mm. One-2 internal dorsal melanophores forming on the forebrain at 10 mm, increasing in number from 13-30 mm but forebrain never entirely covered. One-2 internal dorsal melanophores on hindbrain at 10 mm but hindbrain pigment remaining 350 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

40 • 35 J:: +" • Cl • + r::: :. •• ~ 30 .').+. • + >- "t:J •• • • \.+ 0 m 25 •• " ••••• +..+ • ?!< •••• +~. - .=+- ~-t+. gj •• .,.••+1 a. 20 ~ CIl ...... a..C\I • ~ ~+ i 15 • •+. +

1 0 +--,--,---.---.---r--.-.-.-,--,----r---r---r---r---r---r---r---r---,----,r-~• 2 4 6 8 10 12 14 16 18 20 22 24 Body Length (mm)

D. balteatus + G. gymnoptera I

Figure 19. Dipterygonotus baLteatus and Gymnocaesio gymnoptera. Length of pelvic spine (as a percent of body length) in relation to body length.

sparse and internal. Two melanophores on lower jaw at 10 mm; melanophores on premaxillae at 12 mm and scattered over snout at 18 mm. One internal melanophore at Op spine base and one at mid spine at 7.0 mm (Fig. 20A); three melanophores just below and parallel with Op spine at 23 mm (Fig. 2IA); scattered melanophores around Op spine in largest specimen (Fig. 2IB). One to several melanophores in Dsp2 chevron at 7.0 mm and in Dsp3-5 chevron and associated membrane from 14 mm. In the largest specimen pigment distally on D fin membrane from spine 1 to ray 4 (Fig. 2IB). Distal pigment on P2 ray 1 and associated membrane at 7.0 mm; absent at 20 mm. Dorsolateral band of pigment originating under D rays extending to C peduncle at 23 mm (Fig. 21A). At 30 mm, a band of pigment just ventral and parallel to LL; also scattered dorsolateral pigment anteriorly (Fig. 2IB).

Remarks.- The series consists of larger larvae, but without the gaps that are common in series of other species. It should be recalled that we are not certain that the specimens between 7 mm and 10 mm are G. gymnoptera. This uncertainty arises because the body has not yet become particularly slender and because the development of the procurrent C rays is incomplete. However, these smaller specimens came from the same samples as the larger G. gymnoptera, and because they share most of the elements of the pigment pattern of the larger larvae, it is likely these smaller larvae are G. gymnoptera. Nearly all the available specimens came from a small number of Dana stations in the Indian Ocean off Sumatra. This makes our description especially susceptible to artifacts arising from fading of pigment and limited geographical range. READER AND LEIS: CAESIONINE LUTJANID LARVAE 351

Figure 20. Larvae of Gymnocaesio gymnoptera from eastern Indian Ocean, south west of Sumalra. Scale bars = 1 mm. A. 7.0 mm (AMS 1.34606-001). Identification of this specimen is tentative, !;ee text. B. 11.3 mm (AMS 1.34727-003). C. 17.5 mm (AMS 1.34607-004).

Genus Pterocaesio Pterocaesio chrysozona (Cuvier, 1830)-46 specimens, 3.0-18.9 mm Figures 22-27, Table 8

Material Examined.-Phi1ippine Sea, AMS 1.31538-002: western Coral Sea, AMS 1.26092-003, 1.26150-008, 1.30533-002, 1.30534-002, 1.32556-002, 1.33831-001: western Tasman Sea, AM:S 1.26057-003,1.26114-006, 1.26163-002, 1.26192-003, 1.31382-002: eastern Indian Ocean over northwest continental shelf of Australia, AMS 1.32153-002, 1.33814-001, 1.33815-001, 1.33816-001, 352 BULLETIN OF MARINE SCIENCE. VOL. 59. NO.2. 1996

Figure 21. Larvae of Gymnocaesio gymnoptera. Both specimens are fully scaled, but scales shown only along lateral line. Scale bars = I rom. A. 23.4 rom (AMS 1.34606-001) from eastern Indian Ocean, west of Sumatra. B. 30.1 mm from Jacquinot Bay, New Britain (CSIRO L77). Myomeres not shown as they are obscured by scales.

1.33817-001,1.33818-001,1.33819-001,1.33830-001, 1.33832-001, 1.33833-001, 1.33834-001,1.33835- 001,1.33836-001,1.33837-001,1.33838-001, 1.33839-001, 1.33840-001, 1.33841-001: Andaman Sea, AMS 1.31997-001. Diagnosis.-MoRPHOLOGY. Fin spines long (MRL of Dsp2 49% BL shortly after flexion, Fig. 26) with medium to course serrae on leading and trailing edges; Dsp2 with two leading edges; up to 12 spines on PI border; up to 3 10; 0 Sb; PAsp moderately long, reaching to or beyond pectoral base (MRL 12% BL) (Fig. 25) and weakly serrate in larvae from 10-13 mm; D X-XI,14-16, A m,ll-13. PIGMENT.Initially, four melanophores on the ventral midline of the tail, from just posterior to the anus to the caudal fin anlage (Fig. 22A, B); just after flexion (4.3 mm) a fifth ventral melanophore appearing on the tail (Fig. 23A); no pigment dorsally on trunk or tail until after flexion (4.8 mm); Dsp2 and P2sp with chevron pigment in least developed larva, D fin pigment limited to Dsp2 in all but one larva; a melanophore each anterior and posterior to CS to 4 mm, thereafter, only anterior one present disappearing at 6.9 mm; melanophore on preanal median finfold to 4.8 mm. Description.-MoRPHOLOGY. Body depth moderate to deep (27-42% BL). Head large (32-41 % BL); snout (10-16% BL) equal to or shorter than maxilla (10- 18% BL); maxilla never reaching beyond anterior edge of pupil. Two larval teeth present on premaxillary tip at 3.0 mm; at 5.4 mm teeth present on dentary; no canines present. Head spines smooth except for small serrae on edges of PAsp in some larvae from 10-13 mm and on spines immediately adjacent to PAsp in larvae> 17 mm. PAsp (MRL 12% BL, Fig. 25) reaching to beyond PI base in preflexion larvae (Fig. 22A, B), to PI base in all other larvae except largest in which it falls short. Preopercular spines large, forming early, the least developed READER AND LEIS: CAESIONINE LUTJANID LARVAE 353 specimen (3.0 mm) with PUI 1; PLI 3; PUO 1; PLO 3. Number of preopercular spines increasing to a maximum of PDI 6 and PLI 9 at 17 mm; PUO 9 at 19 mm; PLO 10 at 17 mm; all spines except PUI present in the largest specimen. [0 1 at 3.4 mm; 3 at I7 mm; 2 at 18 mm. Sb 0; Scl 1-2 at 3 mm; 3 at 4.3 mm; 3- 4 from 11 mm. Ptd 1 in preflexion larvae; 1-2 after flexion; 3 at 11 mm; 6 at 19 mm. Ptv 1 at 4.1 mm; absent in the largest specimen. Smooth FS at 3.0 mm; first serration at 4.1 mm; 11 serrae at 17 mm; smooth at 18 mm. Low F from 17 mm. Pe bump at 4.3 mm, forming a short, low ridge by 10 mm; still present in largest specimen. Notochord flexion occurring at 4.1 to 4.8 mm. Three D sp, and Pzsp present in least developed specimen. All fins except PI and procurrent Crays complete at 5.4 mm; procurrent C rays complete at 8.0 mm and PI complete at 9.2 mm. Dsp2 (16-49% BL) always the longest fin spine with Pzsp (11-35% BL) second longest. Fin spines with moderate to coarse serrae. Leading edge ofDspl- 5 and Aspl-2 and trailing edges of all Dsp and Aspl-3 serrate, remaining Dsp leading edges smooth. Leading edge of Asp3 serrate by 9 mm. All fin spine serrations weakening considerably from 16 mm. Lateral-line incipient scales first seen at 17 mm. PIGMENT.Four melanophores on ventral midline of tail until flexion; from jUlst posterior to anus to C anlage. After flexion melanophores at A fin base and peduncle increasing in number. In largest specimen melanophores occuring internally and externally at the base of each D and A fin element, and a line of small melanophores extending posteriorly along anterior 2!J of caudal peduncle ventrally. A larger ventral melanophore on the peduncle just anterior to C fin from postflexion to largest larva. Following flexion one melanophore at base of last 0 fin- ray. From 7.3 mm melanophores at posterior D ray bases increasing in number; three larvae about 9 mm with a melanophore at each D ray base, but most larvae with melanophores at bases of Drays 3-7 and 11-12. Dorsally on peduncle, 1- 2 melanophores at 6.3 mm; at 7.3 mm melanophores extending more than half way along peduncle from D; at 14 mm dorsal pigment on peduncle reaching just short of C. One melanophore at base of lower 2-3 C rays from flexion to 17 mm, thereafter pigment spreading over all upper and 7 lower C rays. In early postflexion larvae. an internal melanophore appearing on dorsal surface of urostyle (Fig. 23A); from 8.7 mm a cluster of external and internal melanophores occuring over urostyle. In least developed larva a melanophore each anterior and posterior to CS (Fig. 22A); during flexion posterior melanophore disappearing; anterior melanophore remaining to 6.9 mm. A small melanophore on preanal median finfold in the least developed larva; disappearing soon after flexion (4.8 mm). An internal melanophore on each side of hindbrain posteriorly, in least developed larva; one external melanophore on each midbrain hemisphere from 4.8 mm; four midbrain melanophores at 6.0 mm. At 7.8 mm, two external melanophores appearing on forebrain. Melanophores increasing in number to a solid covering over the fore-, mid- and hindbrain by 18 mm. From about 6 mm pigment sometimes occuring on snout dorsally, on premaxillae, on dentary, and under chin; all this pigment present in largest specimen. One internal melanophore at opercular spine base at 6.1 mm, thereafter melanophores around Op spine increasing to four at 18 mm. Four melanophores lining orbit of right eye posteriorly in largest specimen. Pigment in Dsp2 chevron and adjacent membrane and in Pz chevron and adjacent ray in the least developed larva; in postflexion larvae melanophores spreading to Pz ray 2 and onto ray 3 at 8-18 mm. Remarks.-The series has a gap between about 14 and 17.5 mm, which is much smaller than the gaps in the series of most other species. Size-specific lengths of the head and fin spines are much more variable in the two Pterocaesio spp. 354 BULLETIN OF MARINE SCIENCE. 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Figure 22. Larvae of Pterocaesio chrysozona from the eastern Indian Ocean over the North West Continental Shelf of Australia. Scale bar = 1 nun. A. 3.0 nun (AMS 1.338]4-(01). B. 3.4 mm (AMS 1.338]4-001). C. 4.4 mm (AMS 1.33817-001). READER AND LEIS: CAESIONINE LUTJANID LARVAE 357

Figure 23. Larvae of Pterocaesio chrysozona from the eastern Indian Ocean over the North \Vest Continental Shelf of Australia. Scale bars = ] mm. A. 4.3 mm CAMS 1.338]6-00]). B. 6.2 mm CAMS 1.33830-00]). described here than in the other caesionines (Figs. 25-27). It is unlikely that this is the result of our mistakenly including other species in our putative series of P. chrysozona and P. tile, especially the latter because of its unique fin meristics. However, it should be remembered that P. chrysozona is the least distinctive caesionine larva we were able to identify, and is easily confused with other species (see section on D. balteatus, above). Therefore, if any of the distributional or meristic data we used to make our identification of P. chrysozona were incorrect, then misidentifications were possible. It is also possible that this intraspeci fic variability is real and a characteristic of the genus Pterocaesio, but we must await the identification and description of the larvae of more Pterocaesio species to test this idea. Much of the description of the larval development of P. chrysozona applies to the development of other Caesio and Pterocaesio spp. larvae we were only alble 358 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2. 1996

Figure 24. Larvae of Pterocaesio chrysozona from the eastern Indian Ocean over the North West Continental Shelf of Australia. Scale bars = 1 mm. A. 8.0 mm (AMS 1.33833-001). B. 11,7 mm (AMS 1.33837-001). C. 17.6 mm (AMS 1.33840-001). Note incipient scales anteriorly on lateral line and serrations on outer preopercular spines. to split into species groups. Much more work remains to be done on the alpha- level taxonomy of the larvae of these two genera. Pterocaesio tile (Cuvier, 1830)-33 specimens, 3.0-16.5 mm Figures 25-29, Table 9 Material Examined.-East China Sea, AMS 1.31801-002, 1.31992-001, 1.32037-002, 1.32089-004, 1.32090-003,1.32094-002: Philippine Sea, AMS 1.31548-001, 1.31549-001, 1.31784-001, 1.31786-002, READER AND LEIS: CAESIONINE LUTJANID LARVAE 359

13 • 12 • • ~ • • • C> 11 •+ + • -t: • • • ~ + 10 • >- • +. + • "'C • 0 + + + • III 9 + • + .-+ •• ;fi + •.+ • ••• • +' + •• • gj 8 + Q. ++ + + • 7 • ~ • ++ + • 6 +

5 2 4 6 8 10 12 14 16 18 20 Body Length (mm)

• P. chrysozona + P. tile

Figure 25. Pterocaesio spp. Length of preopercular angle spine (as a percent of body length) in relation to body length.

1.31822-001: western Coral Sea, AMS 1.24509-003, 1.24516-004, 1.24518-004, 1.24518-005, 1.24519- 006,1.24953-007,1.24954-008,1.24957-004, 1.24958-002, 1.25006-001: Andaman Sea, AMS 1.31988- 001: Mozambique Channel, AMS 1.33120-005, 1.33122-002.

Diagnosis.-MoRPHOLOGY. Spines of all fins long (MRL of Dsp2 37% BL shortly after flexion, Fig. 26) with serrate leading edges and fine comb-like serrae (present also in C. cuning) on trailing edges of P2, Aspl-2 and Dspl-6 (remaining Dsp and Asp3 smooth); single leading edges on D and A spines; PI border with up to 8 spines; 10 1; Sb 0; PAsp with serrae from 5-6 mm, moderately long, reaching to or beyond PI base except in largest larva (MRL 11% BL, Fig. 25); D X- XII,19-22; A III,13. PIGMENT.Initially three melanophores on ventral midline of tail, from just posterior to A anlage to the fin fold near the C anlage; no pigment dorsally on trunk or tail (except in one specimen) until 6 mm; Dsp2 and P2 ray 1 with distal pigment from 5 mm; melanophore inconsistently present at CS from 4.1 mm to 7.7 mm; no pigment on preanal median finfold.

Description.-MoRPHOLOGY. Body depth moderate, initially 35% BL, becoming slender with size (22% BL at 16 mm). Head moderate to large (27-45% BL); snout (7-15% BL) shorter than or equal to maxilla (10-17% BL); maxilla reaching beyond anterior edge of eye, but not to pupil. Teeth present on the premaxillary tip and dentary in preftexion larvae; no canines present. Head spines smooth except PAsp which has small serrae at 5-6 mm; these beginning to reduce from 10 mm but still present at 16 mm. PAsp reaches at least PI base to 10 mm (MRL 11% BL, Fig. 25) thereafter falling short of PI base. Preopercular spines large, forming early; least developed specimen with PUll, PLI 2, PUO 1, PLO 3. 360 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996 50 - 45 ..c. - C) 40 - -r:: - - ~ >- 35 "C 0 -- •• + ++ - • m 30 - ••• + - ?fl. + -+- " -- - + ~ 25 + + + N - a. • ++ - --- - + ~ 20 ~ - + 15 - ++ 10 2 4 6 8 10 12 14 16 18 20 Body Length (mm)

I - P. chrysozona + P. tile

Figure 26. Pterocaesio spp. Length of dorsal-fin spine 2 (as a percent of body length) in relation to body length.

Number of spines increasing with size to maximum of PUI 4; PLI 6; PLO 7, at 16 mm; PUO remaining constant at one. 10 1 from 5.0 mm to largest specimen. No Sb spine throughout development. Scl 1 at 3.4 mm; 2 at 4.7 mm; 3 at 6.5 mm to the largest specimen. Ptd 1 at 3.8 mm; 1-2 at 5.7 mm to the largest specimen. Ptv 1 at 5.7 mm to the largest specimen. Smooth FS at 4.1 mm, 3 serrae at 5.8 mm; 5 serrae at 6.5 mm; smooth at 16.5 mm. Pe at 7.7 mm; F at 8 mm; both Pe and F present in the largest specimen. Notochord flexion occuring at 4.9-5.7 mm (Fig. 28B, C). Dspl-2 and P2sp present in least developed specimen; all fin spines except Asp3 ossified at 7.1 mm; Asp3 ossified only in the largest specimen. All 0, A, C and P2 fin elements present at 5.7 mm; PI and procurrent C rays incomplete at 10 mm, both complete at <16 mm. Dsp2 (14-

37% BL) slightly longer or equal to P2sp, (11-37% BL, Fig. 27) Dspl-6 with fine serrae on leading and trailing edges soon after formation; serrae on trailing edges comb-like; remaining 0 spines smooth. Aspl-2 with fine comb-like serrae

on trailing edges; Asp3 smooth upon ossification. P2 spines with fine comb-like serrae. No scales present in any specimen. PIGMENT. In preflexion larvae three melanophores ventrally on tail, from mid tail to notochord tip (Fig. 28A). After flexion melanophores arranged as follows: one just posterior to A base; one half way along peduncle; one at lower C ray bases; this ventral pigment retained to largest specimen except for a 10 mm larva with pigment at base of A rays 6-13 (Fig. 29B). External dorsal melanophore present on notochord in one larva at 4.1 mm (Fig. 28A); at 5.8 mm two melanophores dorsally on peduncle; at 8,6 mm melanophores on anterior half of peduncle dorsally; at 10 mm melanophores from Dsp3 posteriorly along 0 base and peduncle to C. Throughout series one to three melanophores found over lower 2- READER AND LEIS: CAESIONINE LUTJANID LARVAE 361

40 + 35 ~ - +++ *+ ++ •..OJ - + r:: + !I!+ + + (I) 30 + ....I ~ -+ >- + + - "C +. - •• 0 OJ 25 - - --- "* .. - .. +

- P. chrysozona + P. tile Figure 27. Pterocaesio spp. Length of pelvic spine (as a percent of body length) in relation to body length.

4 C rays. The dorsal melanophore on notochord of 4.1 mm larva probably becoming the internal pigment spot on dorsal surface of urostyle. In postflexion larvae, urostyle pigment increasing in area both internally and externally, and remaining in largest specimen. Six of 15 larvae between 4.1 mm and 7.7 mm with a melanophore anterior to the CS. At 4.9 mm 2 dorsal melanophores on midbrain and two internal melanophores between junction of mid- and hindbrai.n; at 10 mm midbrain (externally) and hindbrain (internally) completely pigmented, and four dorsal melanophores on forebrain (Fig. 29B); the forebrain almost completely pigmented in largest specimen. Melanophores on premaxillae and dentary tip from 6-7 mm. Pigment present on snout dorsally in two specimens at 8.6 mm and 10 mm and under chin at 10 mm (Fig. 29B). An internal melanophore at Op spine base at 7.7-10.3 mm; two external melanophores on Op spine base in largest specimen. Dsp2 chevron pigment present in six of 19 larvae from flexion. Pigment on P2sp and ray 1 distally first seen at 4.7 mm, again at 7.0 mm; thereafter in 3 of 10 specimens to 16 mm. Remarks.-The series contains only one larva larger than 10 mm. The gap between 10 and 16 mm is readily bridged via the unique fin meristics of P. tile (Table 1) and its distinctive fin-spine ornamentation. However, the variability in BL-specific spine lengths (Figs. 25-27) and variability in pigment (above) sug- gests the possibility that P. tile as presently conceived contains one or more cryptic species. Geographic variation in this very wide-spread species is also a possibility. Unfortunately, our material is inadequate to assess either of these possibilities. From examination of the condition of the specimens, it appears th.at some of the apparent variability in BL-specific spine length relationships is due 362 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

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Figure 28. Larvae of Pterocaesio tile from the Great Barrier Reef, near Lizard Island. Scale bars = 1 mm. A. 4.1 mm (AMS 1.24953-007). B. 4.9 mm (AMS 1.24957-004). C. 5.4 mm (AMS 1.24509- 003). READER AND LEIS: CAESIONINE LUTJANID LARVAE 365

Figure 29. Larvae of Pterocaesio tile. Note serrate PAsp. Scale bars = ] mm. A. 7.7 mm from Great Barrier Reef, near Lizard Island. (AMS 1.24519-006). B. ]0.3 mm from Great Barrier Reef, near Lizard Island. (AMS 1.245]8-005). C. ]6.5 mm from the western Coral Sea. (AMS 1.25006-001). 366 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996

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0 U= >,.c 31000'" 0\ '8~ ,...,NII'\ IU 0\0\0..0 Z CQ,2 ..•..• ~ 368 BULLETIN OF MARINE SCIENCE, VOL. 59, NO.2, 1996 to shrinkage in BL of smaller specimens (see Leis and Lee, 1994). A high proportion of P. tile specimens came from mid-water trawls not taken for study of fish larvae. Larvae are bycatch in such samples and as a result they are likely to be damaged or to have suffered shrinkage due to poor handling and delay in fixation.

ACKNOWLEDGMENTS

This study could not have proceeded without the generous cooperation of many people who made available the collections of larval caesionines in their care, made room in their laboratories, and in many cases deposited the larvae at AMS or other accessible archival institutions. In particular, we would like to acknowledge L. Beckley, E. Bertelsen, B. B. Collette, P. J. Doherty, S. English, A. Graham, K. E. Hartel, R. F. Hartwick, H. F. Hausfeld, Y. Iwatsuki, G. D. Johnson, J.-I. Kojima, P. Last, R. J. Lavenberg, M. Miya, K. Matsuura, G. E. McGowan, N. E. Milward, J. Nielsen, M. aki- yama, T. Ozawa, J. R. Paxton, B. Rachod, W. J. Richards, T. Saruwatari, P. H. Schalk, D. McB. Williams, and P. C. Young. D. J. Bray, S. Bullock, K. Lee, and T. Trnski ably assisted in the lab. T. Goh assisted with data entry and production of the typescript. B. C. Mundy provided valuable con- structive criticism of the manuscript. Thanks also to D. F. Hoese for allowing S.E.R. the time to complete this manuscript. Many larvae utilized in this study were collected with support provided by Australian Marine Science and Technologies Grants 80/2016 and 83/1357 to J. M. Leis and B. Gold- man, and an Australian Research Council (ARC) Grant 8610873 to J. H. Choat and J. M. Leis. This project was supported by ARC grant A19031159, and by the Australian Museum. K. E. Carpenter provided much useful information on caesionines. Our very great thanks to all.

LITERATURE CITED

Allen, G. A. and R. Swainston. 1988. The marine fishes of north-western Australia. Western Austra- lian Museum. 201 p. Anderson, W. G., Jr. 1987. Systematics of the fishes of the family Lutjanidae (Perciformes: Percoidei), the snappers. Pages 1-28 in J. J. Polovina and S. Ralston, eds. Tropical snappers and groupers: biology and fisheries management. Westview Press, Boulder Colorado. 659 p. Bell, L. J. and P. L. Colin. 1986. Mass spawning of Caesio teres (Pisces: Caesionidae) at Enewetok Atoll, Marshall Islands. Env. BioI. of Fishes. 15: 69-74. Carpenter, K. E. 1987. Revision of the Indo-Pacific fish family Caesionidae (Lutjanoidea), with descriptions of five new species. Indo-Pacific Fishes No. 15. 56 p. ---. 1988. Fusilier fishes of the world. FAO species catalogue No. 125, Vol. 8.75 p. ---. 1990. A phylogenetic analysis of the Caesionidae (Perciformes: Lutjanoidea). Copeia 1990: 692-717. Fourmanoir, P. 1976. Formes post-Iarvaires et juveniles de poissons cotiers pris au chalut pelagique dans Ie sudouest Pacifique. Cah. Pacif. 19: 47-88. Hamner, W. M., M. S. Jones, J. H. Carleton, I. R. Hauri and D. McB. Williams. 1988. Zooplankton, planktivorous fish, and water currents on a windward reef face: Great Barrier Reef, Australia. Bull. Mar. Sci. 42: 459-479. Johnson, G. D. 1980. The limits and relationships of the Lutjanidae and associated families. Bull. Scripps Instit. Oceanogr., Univ. Calif., San Diego 24: 1-114. ---. 1984. Percoidei: development and relationships. Pages 464-498 in H. G. Moser, W. J. Rich- ards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr. and S. L. Richardson, eds. Ontogeny and systematics of fishes. Amer. Soc. Ichthyol. Herpetol. Spec. Pub. 1. 760 p. ---. 1993. Percomorph phylogeny: progress and problems. Bull. Mar. Sci. 52: 3-28. Kojima, J. I. 1988. Lutjanidae. Pages 511-517 in M. Okiyama, ed. An atlas of the early stage fishes in Japan. Tokai Univ. Press, Tokyo. 1154 p. (In Japanese) Leis, J. M. 1987. Review of the early life history of tropical groupers (Serranidae) and snappers (Lutjanidae). Pages 189-237 in J. J. Polovina and S. Ralston, eds. Tropical snappers and groupers: biology and fisheries management. Westview Press, Boulder, Colorado. 659 p. --- and D. J. Bray. 1995. Larval development in the lutjanid subfamily Paradicichthyinae (Pisces): the genera Symphorus and Symphorichthys. Bull. Mar. Sci. 56: 418-433. --- and K. Lee. 1994. Larval development in the lutjanid subfamily Etelinae (Pisces): the genera Aphareus, Aprion, Etelis and Pristipomoides. Bull. Mar. Sci. 55: 46-125. --- and D. S. Rennis. 1983. The larvae of Indo-Pacific coral reef fishes. New South Wales Univ. Press, Sydney and Univ. Hawaii Press, Honolulu. 269 p. --- and T. Trnski. 1989. The larvae of Indo-Pacific shorefishes. New South Wales Univ. Press, Sydney and Univ. Hawaii Press, Honolulu. 371 p. READERANDLEIS:CAESIONINELUTJANlDLARVAE 369

Leviton, A. E., R. H. Gibbs, Jr., E. Heal and C. E. Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985: 802-832. Randall, J. E., G. R. Allen and R. C. Steene. 1990. Fishes of the Great Barrier Reef and Coral Sea. Crawford House Press, Bathurst, Australia. 507 p. Springer, V. G. 1982. Pacific plate biogeography, with special reference to shorefishes. Smithsonian Contrib. Zool. 367: 1-182. Thresher, R. E. 1984. Reproduction in reef fishes. T.F.H. Publications, Neptune City, New Jersey. 399 p. Yokoyama, K., Y. Kamei, M. Toda, K. Hirano and Y. Iwatsuki 1994. Reproductive behavior, eggs and larvae of a caesionine fish, Pterocaesio digramma, observed in an aquarium. Japn. J. Ichthyol. 41: 261-274. ---, Y. Kamei, M. Toda, K. Hirano and Y. Iwatsuki. 1995. Reproductive behavior, eggs and larvae of a caesionine fish, Caesio caerulaurea, observed in an aquarium. Japn. J. Ichthyol. 42: 157- 164.

DATEACCEPTED: June 12, 1995.

ADDRESS: Fish Section, Australian Museum, 6-8 College St., Sydney, NSW, 2000, Australia.