Larval Development in the Lutjanid Subfamily Etelinae (Pisces): the Genera <I>Aphareus, Aprion, Etelis</I> and <I

BULLETIN OF MARINE SCIENCE, 55(1): 46-125. 1994 CORAL REEF PAPER

LARVAL DEVELOPMENT IN THE LUTJANID SUBFAMILY ETELINAE (PISCES): THE GENERA APHAREUS, APR/ON, ETEL/S AND PR/STIPOMOIDES

Jeffrey M. Leis and Krysia Lee

ABSTRACT Larval development in the lutjanid subfamily Etelinae is described and illustrated from field specimens captured with plankton nets and midwater trawls in the Atlantic, Indian and Pacific Oceans. Series were assembled using fin-spine morphology, scale counts, gill-raker counts, and pigment patterns. Identifications were confirmed from adult characters developed in the largest specimens and from the distributions of the adults. Larvae of the Etelinae share with larvae of other lutjanids the presence of a spine on the dorsal postcleithrum, early forming, very elongate spines of dorsal and pelvic fins, and even longer pelvic-fin soft rays. They are further characterized by fin-ray counts (D X,IO; A 111,8),absence of external ornamentation on fin spines (except Aprion where weak, trailing-edge serrations are present). early forming pigment on the midbrain posteriorly, and early formation of both third anal- fin spine and scales. Aprion virescens is characterized by serrate fin spines, an especially long second spine in the dorsal fin, pigment series on the longest dorsal-fin spines, pigment on the urostyle and absence of dorsal pigment on trunk and tail. A. virescens is pelagic until at least 18 mm. Etelis spp. are characterized by a long, fragile second spine in the dorsal fin, fin spines lacking internal structure, pigment series on the pelvic-fin spine and longest dorsal- fin spines, absence of dorsal pigment on trunk and tail, ventral limited pigment on tail that disappears early, pigment on the forebrain that forms early, and absence of urostyle pigment. We lacked specimens between 8.6 and 13.7 mm. Three Indo-Pacific types of Etelis larvae unidentified to species and distinguished by minor pigment and preopercular spine differences are recognized in specimens <8.6 mm. Specimens> 13.7 mm are identified as E. carbunculus. E. coruscans (Indo-Pacific) and E. oculatus (Atlantic), primarily on the basis of counts of scales and gill rakers and of geographical distributions. Etelis spp. are pelagic to at least 51 rom. Aphareus and Pristipomoides larvae are virtually indistinguishable and are characterized by a moderate-sized second spine in the dorsal fin (often shorter than pelvic-fin spine), fin spines with distinctive internal structure, a pigment cluster at the base of both anal fin and dorsal fin posteriorly, urostyle pigment at some period during development and lack of pigment series on the dorsal-fin spines. The Indo-Pacific A. rutilans, P. auricilla, P. filamentosus, P. multidens and/or typus, P. sieboldii, and two unidentified members of Aphareus or Pristipomoides (with incomplete series) and the Atlantic P. aquilonaris and what is possibly P. freemani are differentiated by lengths of dorsal-fin spines, minor differences in pigmentation, numbers of lateral-line scales and gill rakers, and geographical distribution. Aphareus rutilans also has a shorter ascending process of the premaxilla than do the Pristipomoides spp. Aphareus and Pristipomoides spp. are pelagic to at least 54 rom. Two types of larvae from the western Pacific with incomplete series (maximum size, 5.7 rom) of a total of five specimens cannot be identified to genus (they are not Randallichthys). They are characterized by deep bodies, relatively short dorsal fin spines, fin spines that lack ornamentation or internal structure, very light pigment and small size at flexion. The putative species are questionably separated by minor pigment differences, size at flexion, body depth, and fin-spine length. Evidence from larvae supports the distinctiveness of Etelis and Aprion and indicates that Aphareus and Pristipomoides are very closely related. The unidentified larvae from the western Pacific are important, for if they prove to be P. argyrogrammicus and/or P. zonatus, they will indicate that these two species are probably not closely related to the other species of Pristipomoides.

Lutjanids of the Subfamily Etelinae (sensu Johnson, 1980) are medium to large carnivorous fishes of tropical to warm temperate waters of both the Atlantic and

46 LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 47

Indo-Pacific. Aprion (one species) and Aphareus (two species) inhabit relatively shallow coral reefs, whereas Etelis (four species), Pristipomoides (11 species) and Randallichthys (one species) live in greater depths (to 550 m) mostly on rocky bottoms (Allen, 1985). Pristipomoides species may be semi-pelagic. Many eteline species have extremely wide distributions. Aphareus furca, for example is found from southern Africa across the Indian and Pacific Oceans to islands off the Pacific coast of central America (Allen, 1985; Rosenblatt et aI., 1972). As another example, Pristipomoides sieboldii is the only lutjanid to occur in the Atlantic, Indian and Pacific Oceans (Anderson 1987). Many eteline lutjanids are important to commercial and artesinal fisheries, and are caught by methods ranging from hook and line to bottom trawl (Allen, 1985). Jobfish seems to be the most widely used common name for species of this subfamily (Allen, 1985). Lutjanid larvae are striking, as they have very elongate dorsal- and pelvic-fin spines, long pelvic-fin rays, "flag-like" pigment on these fins distally, and extensive spination on the head (Leis, 1987). Unfortunately, larval development has been described for only a few of the many species (see review by Leis, 1987, and also Potthoff et aI., 1988; akiyama, 1988, 1991; Iwatsuki et aI., 1991). Knowledge of the early life history of eteline species is particularly limited. Spawning mode is known for only two eteline lutjanids: Pristipomoides sieboldii and P. filamentosus spawn pelagic eggs, as do alI other lutjanids for which spawning mode is known (Leis, 1987). Larvae are virtually undescribed. Fourmanoir (1976) mentioned larvae of Etelis oculatus (obviously mis-identified as he was reporting on Indo-Pacific material; 29-59 mm), E. carbunculus (44 mm) and Pristipomoides multidens (22-35 mm), but did not describe them other than to give standard lengths and some meristic values. Leis (1987) illustrated a 7.1 mm larva of Aprion virescens (the illustration was reproduced by Allen, 1985), and provided a table comparing the characteristics of the larvae of lutjanid subfamilies. The larvae of eteline lutjanids are distributed mostly seaward of the edge of the continental shelf, and offshore of oceanic islands (Leis, 1987). This is in contrast to most larvae of the subfamilies Caesioninae and Lutjaninae which are found primarily over the continental shelf. This means that the collections by the DANA (held at ZMUC), by Japanese cruises in the oceanic western Pacific, and by Hawaiian scientists studying midwater organisms were especially valuable sources of eteline larvae for study. In spite of our best efforts to obtain larval etelines from collections held in North America, Europe, Japan, Hawaii and Australia, many series are incomplete or contain gaps, a number of series are incompletely identified, and several taxa remain unidentified. This lack of material has been a major frustration during this study. However, it is unlikely more material will become available soon. We include here descriptions of all the series we were able to assemble, complete or not, fully identified or not, so future investigators will not have to duplicate our work, and so the available material (and its location) of the taxa that require further work will be documented. Further, it was essential to document the entire range of ontogenetic diversity in the subfamily to allow later use of this ontogenetic information in a planned assessment of the relationships of the Lutjanidae. It is the purpose of this paper to describe the larval development of as many eteline lutjanid species as possible. Our original intent was to describe only Indo- Pacific species. We were aware that a number of workers had more-or-Iess complete series of an Atlantic Pristipomoides (including L. A. Collins, A. W. Kendall, Jr., K. C. Lindeman, and W. J. Richards), and we were content to leave the description of this taxon to them. When these workers became aware of our work on Indo-Pacific species, all very generously turned over to us their notes or spec- 48 BULLETIN OF MAR[NE SCIENCE, VOL. 55, NO.1, [994 imens and encouraged us to include the Atlantic taxa in our study. Thus the present paper is able to include descriptions of varying degrees of completeness of larval development of 16 species (some unidentified beyond the genus or subfamily level) including four identified genera. Only the rare Randallichthys amongst the five genera of the subfamily could not be included due to lack of material. The larval development of these species provides information that will shed light on the relationships of the subfamily and within it. This information will be mentioned only briefly here, as it will be more fully analyzed in a separate paper. This paper is the first in a series which will describe the larval development of lutjanid fishes. It is planned to produce a paper on each of the remaining subfamilies (Apsilinae, Lutjaninae, Paradicichthyinae, and Caesioninae) in the same format as this one.

MATERIALS AND METHODS

When establishing identifications, we assumed there are no valid taxa in the subfamily other than those recognized by Johnson (1980), Anderson (1987) and Allen (1985): thcse are listed in Table 1. The known species are moderate to large in size, widely distributed and of commercial importance, so the assumption is reasonable. However, they occur in difficult-to-sample, relatively deep habitats, and have their greatest diversity in the Indo-Pacific, the part of the world ocean where ichthyological exploration is most incomplete. Further, the most recently discovered species in the subfamily was described by W. D. Anderson, Jr. in 1981, so some undescribed taxa may remain. Definitions, measurements and abbreviations of them follow Leis and Trnski (1989) with the exception that the eye diameter is of the orbit, not the pigmented area of the eye. Abbreviations used are: A-anal fin; BD-body depth; BL-body length; D--dorsal fin; LL--lateralline; MRL-maximum relative length; P,-pectoral fin; P2-pelvic fin; sp--spine (used in conjunction with abbreviations of fins or bones: e.g., Dsp2 is the second spine of the dorsal fin). Abbreviations for the spines of the head are generally based on the bone from which the spine or ridge arises, and are adapted from those of Johnson (1984): F-low, anterior ridge on the frontal; FS-supraorbital ridge of the frontal bone; P I-inner border of the preopercle; P O--outer border of the preopercle (U- upper, and L--lower, used in conjunction with POor P Trefer to the limbs of the inner and outer preopercular border: e.g., PUO is the outer border of the upper limb of the preopercle); PAsp--large spine at the angle of the preopercle; Op--opercle; Sb-subopercle; Io-interopercle; Pt-posttemporal; Scl-supracleithrum; PcI-postcleithrum; Pe-pterotic. A notched dorsal fin is one in which the ultimate spine is longer than the penultimate. Gill-raker counts are of the lower limb of the first arch, do not include the raker at the angle, and were made only on larvae >8.0 mm (except that Table I gives counts of the full arch). 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 tubes of the lateral line scales were not fully formed, or they could not be discerned, particularly posteriorly. Forming scales were difficult to count, and some scales were frequently missing, so many of the counts are necessarily approximate. For meristic values, non-integer counts indicate bilateral asymmetry, and represent mean values. 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 still retain temporary specializations for pelagic life. These are primarily the spination of the head (albeit reduced in the larger specimens) and pigment patterns not found in the bottom-associated juveniles and adults. Larvae were examined under a dissecting microscope. Precision of measurements varied with the magnification used: this varied from 8 to SOx depending on size of the specimen and the structure being measured. Most measurements were made with a digitizing pad/camera lucida system attached to a Wild M5 dissecting microscope. Illustrations were prepared using a Zeiss SR dissecting microscope equipped with an adjustable drawing tube. Material Examlned.-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. In the "Remarks" section of the description of Aprion virescens is a discussion of shrinkage of specimens and its effects on morphometries. This undoubtedly applies to other species as well. Many specimens were damaged and the elongate spines of the dorsal and pelvic fins and rays of the latter were particularly prone to breakage. The pigment associated with these spines and their intervening membranes is frequently missing in part or whole due to damage. This limits the utility LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 49

Table 1. Meristic characters of eteline lutjanids. Compiled from: Kami (1973), Senta and Tan (1975), Anderson et al. (1977), Kyushin et al. (1977, 1982), Johnson (1980), Anderson (1981, 1987), Okamura et al. (1982), Uyeno et al. (1983), Masuda et al. (1984) and W. J. Richards (pcrs. comm.). Pristipo- moides lateral-line scale counts from Kami (1973) are consistently 3-5 higher than those from An- derson (1987), even though both claim to use the same convention for counting. The tabled counts are from Anderson (1987). The four Allantic species may all rarely have 10 dorsal-fin rays.

Fin rays Lateml Supra- line neural Dorsal AnaJ Pectoral Gill rakers scales bones Indo-Pacific species Aphareus A. jurca X,11 m,8 16 6-10 + 16-18 = 22-27 69-75 3 A. rutilans X,10-11 m,8 15-16 16-19 + 32-35 = 49-52 69-75 3 Aprion A. virescens X,ll m,8 16-18 7-8 + 14-16 = 21-24 45-50 2 Etelis E. carbunculus X,11 m,8 15-17 5-8 + 11-14 = 17-22 48-50 3 E. coruscans X,11 m,8 15-16 8-10 + 15-18 = 23-28 47-50 3 E. radiosus X,11 m,8 16 12-15 + 20-22 = 33-36 50-51 3 Pristiopomoides P. argyrogrammicus X,11 m,8 15-16 5-6 + 12-15 = 17-21 59-61 3 P. auricilla X,11 m,8 15-16 9-11 + 18-19 = 28-30 67-7] 3 P. filamentosus X,I] m,8 15-16 7-8 + 15-]7 = 23-25 57-62 3 P. jlavipinnis X,11 m,8 16 6-8 + 16-17 = 22-25 59-62 3 P. multidens X,I] m,8 16 6-8 + 13-15 = 20-22 48-50 3 P. sieboldii X,lI m,8 16-17 9-12 + 21-22 = 30-33 69-71 3 P. typus X,II m,8 16 6-9 + 1]-16 = 18-25 48-50 3 P. zonatus X,11 III,8 16-17 5-8 + 12-]4 = 17-21 63-67 3 Randallichthys R. filamentosus X,ll III,9 16-17 5-9 + 14-]6 = 19-23 48-49 3 Atlantic species Etelis E.oculatus X,ll III,8 15-17 7-]1 + 14-18 = 23-28 47-50 3 Pristipomoides P. aquilonaris X,IO-11 III,7-8 15-]7 7-9 + 16-20 = 24-28 47-52 3 P. freemani X,II-12 m,8 15-17 8-10 + 19-23 = 28-32 47-52 3 P. macrophthalmus X,I] m,8 15-17 6-8 + 13-]7 = 19-25 54-57 3 of characters based on spine length and fin pigment, which is unfortunate, as these are important in distinguishing species within genera. Larvae of only two ete]ine species-A. virescens and P. filamentosus-were found in near-shore plankton samples taken from small boats. Because of the way these samples were handled, they have provided preflexion and flexion-stage specimens in exceptional condition, and this has allowed the descriptions of the preflexion larvae of these two species to be especially detailed. Some larvae, especially those which have long been 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. Most material had been fixed in formalin, but concentration of formalin and buffers used varied widely. By the time they were examined for this study, most specimens were preserved in 70% ethanol. 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 smaller size increments, surplus material was frequently available. In some cases, more than the minimum five per mm increment were examined in detail, but the surplus material was examined for diagnostic characters. Surplus material is not listed in the "Material Examined." Institutional abbreviations follow Leviton, et al. (1985), with the following additions: NMFSU and NMFSM which indicate National Marine Fisheries Service, La Jolla, California, and Miami, Florida, respectively; and MUFS which indicates Miyazaki University Fisheries Science, Miyazaki, Japan. The Literature Cited section does not include citations based solely on the authorities for the genera and species. These are given by Allen (1985) and Anderson (1987). 50 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

IDENTIFICATION No reared material was available, so all identifications are based on the series method. Not all species of the subfamily were identified. Until larvae of all species are identified, there will remain 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 putative series are not mono- specific. Throughout, we have avoided distinguishing specimens based on only a single character (e.g., Pristipomoides filamentosus with and without the cleithral melanophore). A corroborating character was required. This approach has the virtue of not over-splitting, but it could lead to over-lumping. For this reason, it is worth examining the taxa not identified. No Randalliehthys filamentosus larvae were identified, but if any were among the study material, they would have been detected due to their count of A III, 9 (Table 1). Aphareus furea was not identified, but it is possible it is the larval type called Pristipomoides sp. S. Amongst the Indo-Pacific Pristipomoides were three species whose larvae were not identified: P. argyrogrammieus, P. flavipinnis, and P. zonatus. They may be included among the following partial series: Pristipo- moides sp. P,Etelinae sp. 1 and Etelinae sp. 2. More material is required to resolve this. Among the Atlantic Pristipomoides, only one, and possibly two out of three species were identified. Here there is more possibility that the series is not mono- specific. This was looked for carefully, but no evidence of it was found. 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 spines, which are smooth to coarsely serrate, but without recurved spinelets; longest pelvic ray longer than pelvic spine; strong head spination, without serrations on preopercular spines (except some lutjanine and caesionine taxa, and in these the serrations are weak); head spination includes a weak anterior frontal ridge, a supraocular ridge, a weak pterotic ridge, and spines on the inner and outer borders of the preopercle, the opercle, suboperc1e, and interoperc1e; spines are also present on the posttemporal, supracleithrum and dorsal postcleithrum; no supraoccipital crest or spines; no lachrymal spines or serrations; body and head laterally compressed; and a series of ventral melanophores on tail at least in preflexion larvae.

Etelinae.-The fin counts of D X,11 and A 111,8and supraneural formula of the larger specimens (0/0+0/2/1 + 1I, except Aprion which has 010/2/1 + 1/) confirmed that they are members of the subfamily Etelinae. No other lutjanids have the fin- ray counts of etelines, and non-eteline lutjanids have a supraneural formula of 0/0/0+2/1 + 1 (except Dipterygonotus which has 10+0/2/1 + 1/). Eteline larvae are further characterized by a weak supraocular (frontal) ridge without serrations; external ornamentation on fin spines lacking or (in Aprion) consisting of weak serrations only; early development of scales; and early development of dorsal pigment on the midbrain. Larvae of Pristipomoides and Aphareus are extremely similar, and cannot be distinguished at the generic level (Table 2). Individual species can be distinguished with difficulty on the basis of minor differences in morphology and pigment of the sort which otherwise distinguish species within genera. None of the major differences in fin-spine structure found among the other genera are present between Pristipomoides and Aphareus, nor are there major differences in pigment pattern such as the dashed-line of melanophores in the longest dorsal spines of LEIS AND LEE: ETELlNAE LARVAL DEVELOPMENT 51

Etelis or the lack of dorsal pigment in Etelis and Aprion. The only non-pigment characters found so far to distinguish Pristipomoides from Aphareus are the late formation of subopercular spines and the short ascending process of the premaxillae in the latter (but based only on A. rutilans), and these are useful only in larger larvae (Tables 3, 4). Two types of larvae for which a full series was not available could be identified only to "Pristipomoides or Aphareus" because of the similarity between the two genera. For convenience, these are herein referred to as Pristipomoides species P and species S. APHAREUS RUTILANS. This species is very similar to Pristipomoides spp., especially P. sieboldii. A series was assembled and linked together through the smooth fin spines with late-forming internal structure, spine length and pigment- most notably a cluster of melanophores at the posterior end of the dorsal fin and a similar anal-fin cluster, presence of melanophore at cleithral symphysis in smaller specimens, absence of pigment at tip of lower jaw, extensive pigment on the pelvic fins and a pigment cluster concentrated on the ventral side of the urostyle (Table 3). The larger specimens have 70 or more lateral-line scales, and many gill rakers (Fig. 6). The scale count eliminates all etelines except two Pristipo- moides spp. (sieboldii and auricilla) and the two Aphareus spp (Table 1). The high gill-raker count alone (>30 rakers on the lower limb) confirms an identification of A. rutilans (Table I). In addition, the ascending process of the premaxilla is short, which corroborates an identification of Aphareus, because Adult Aphare- us have a short process associated with non-protrusible premaxillae (Johnson, 1980). APR/ON V/RESCENS.A series was assembled, initially from Hawaiian material, and linked together through serrations on the fin spines, spine length and pigment-most notably lack of dorsal pigment on trunk or tail, and the distinctive internal melanophores on the lower jaw (Table 2). The largest specimen has 18 pectoral rays and only two supraneural bones, both of which confirm an identification of Aprion (Table 1). Aprion is monotypic, so this larval series was identified as Aprion virescens. ETELIS. A series of each of six types (three of larvae <8.6 mm and three of larvae> 13.7 mm, see below) was assembled and linked together through the smooth, fragile, fin spines which lack visible internal structure, and through pigment-most notably lack of dorsal pigment on trunk or tail, distinctive series of melanophores in the posterior groove of the pelvic-fin spine and dorsal-fin spine 2, lack of ventral pigment on trunk and tail after the flexion stage, and early development of melanophores on the forebrain (Table 2). Larvae larger than 40 mm have scales on the maxilla, a feature unique to Etelis among adult etelines. In the two largest specimens with intact dorsal-fin spines, the penultimate spine is shorter than the ultimate, also a feature unique to Etelis among adult etelines. Therefore, the larvae were identified as Etelis. Unfortunately, a size gap in the series between 8.6 and 13.7 mm introduces some uncertainty for the smaller specimens, but there is nothing to indicate that larvae from more than one genus are included. Corroboration of the identification comes from several independent characters. Specimens larger than 7 mm have about 50 lateral-line scales: these eliminate Aphareus spp. and Pristipomoides spp. (except typus and multidens). Those with full fin-ray counts have 16 pectoral rays and three supraneural bones which eliminate Aprion, and A III,8 which eliminates Randallichthys. This leaves only Etelis spp., P. typus and P. multidens. Larvae of Pristipomoides spp. (including at least one of the two latter species) have relatively consistent fin-spine structure and pigment pattern, both of which differ from that of the present larvae. This leads to an identification of Etelis. 52 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

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The Atlantic specimens are attributed to the sole Atlantic species of the genus, Etelis oculatus. Gill-raker counts (Table 1) indicate that among the six largest Indo-Pacific specimens are Etelis carbunculus (11-12 rakers, N = 3), and Etelis coruscans (16-17 rakers, N = 3). No specimens are attributable to the third Indo- Pacific species, Etelis radiosus. Unfortunately, because of the size gap, smaller specimens cannot be identified to species. Among the smaller Indo-Pacific specimens are two species of Etelis, the largest specimens of which have scales and fins formed. A third species is probably a member of Etelis based on fin spine morphology and pigment, but the largest specimens lack scales and formed pectoral rays, so this cannot be confirmed. These three taxa are differentiated by pigment and by the number of spines on the outer border of the preopercle. PRISTlPOMOIDES. Series of each of eight types were assembled and linked together primarily through length, internal structure and absence of external ornamentation of the dorsal and pelvic spines, number of lateral-line scales, number of gill rakers, number of preopercular spines and pigmentation, particularly a cluster of melanophores at the posterior end of the dorsal-fin base, and a corresponding anal-fin cluster, a spot generally centered over the urostyle, the presence or absence of melanophores on the bases of dorsal and pectoral fins, and the number of melanophores at the c1eithral and lower jaw symphyses (Table 3). The largest specimens of those series herein considered as P. multidensltypus, P. sieboldii, P. aquilonaris and P. filamentosus were identified as Pristipomoides using the key of Johnson (1980), with the aid of cleared and stained material where necessary. All have A III,8 (eliminates Randallichthys), three supraneurals (eliminates Aprion), long ascending processes on the premaxillae (associated with protrusible premaxillae in adults and which eliminates Aphareus and Randallichthys), unsealed maxillae, and non-incised dorsal fin (both of which eliminate Etelis). In the four other species, the largest individuals in the size series were too small to be identified by the above criteria: they were considered to be Pristipomoides because of their similarity to the other Pristipomoides spp. The characters used to construct the series are summarized in Tables 2 and 3. Indo-Pacific Pristipomoides species (there are eight Indo-Pacific species, Ta- ble 1). PRISTlPOMOIDES AURICILLA. The only specimen identified as this species has 70- 72 lateral-line scales, which eliminates all etelines except the two Aphareus species, P. sieboldii and P. auricil/a. The gill-raker count of 15 at 12 mm eliminates A. rutilans (Fig. 6). The strong internal structure in the fin spines eliminates P. sieboldii, and it appears that this specimen has fewer gill rakers than do similar- sized specimens of P. sieboldii (Fig. 6). The ascending process of the premaxilla is long (Fig. 8), indicating Pristipomoides auricil/a rather than Aphareus furca (Table 3). PRISTlPOMOIDES F/LAMENTOSUS. Fully-scaled specimens have 60-64 lateral-line scales. This count eliminates all but P. argyrogrammicus, P. jlavipinnis and P. filamentosus. The five largest specimens have 20-23 gill rakers, which, while not the final complement, eliminate P. argyrogrammicus. The majority of the initially available specimens came from Hawaii where, of the three species, only P. filamentosus occurs. There was no indication that the non-Hawaiian material consti- tuted a species different from the Hawaiian material. Therefore, this series was identified as P. filamentosus. However, until larvae of P. jlavipinnis are identified, the possibility remains that they are included among the (non-Hawaiian) material described here as P. filamentosus (Table 3). PRISTlPOMOIDES MULTIDENS or TYPus. Fully-scaled specimens have 48-53 lateral- line scales. Only P. typus and P. multidens have this few lateral-line scales (Table LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 57

3). These two species are very similar-adults can only be distinguished with difficulty- and they have broadly overlapping distributions (Allen, 1985). We are unable to determine which species is represented in this series (see "Remarks" under the description of these larvae). PRISTIPOMOIDES SIEBOWl/. Fully-scaled specimens have 69-72 lateral-line scales. This count eliminates all Pristipomoides spp. but sieboldii and auricilla. Aphareus spp. are eliminated by gill-raker counts and length of the ascending process of the premaxilla (Figs. 6, 8). No vomerine or tongue teeth are developed, and gill rakers are apparently not fully developed (Fig. 6), so these characters could not be used. However, most of the specimens were from areas where P. auricilla is rare and P. sieboldii common (Hawaii, J. E. Randall, pers. comm.) or areas where P. auricilLa is not known and P. sieboldii is known (Australian east coast, Allen, 1985, and our unpubl data). Further, 9 of 12 specimens with a full complement of rays in the pectoral fin had 17 rays in both fins (the other 3 had 16). Information provided by W. D. Anderson, Jr. (pers. comm.) and our own counts indicate that P. sieboldii usually (10 of 16 specimens) has 17 pectoral rays while P. auricilla has 15 or 16 (10 specimens, including two listed by Abe, 1960). We conclude, therefore, that the series was composed of P. sieboldii (Table 1). PRISTIPOMOIDES SPECIES S. Fully scaled specimens have 69-73 lateral-line scales, which indicates an identification of Aphareus spp. or one of only two Pristipo- moides spp. (sieboldii and auricilla). Unfortunately, none of the specimens are large enough to provide definitive gill-raker counts (Fig. 6), but the gill-raker counts are too low for these larvae to be Aphareus rutilans, which is, in any case, already identified above. If it is assumed they are not P. sieboldii, which is identified above, this leaves only Aphareus furca and Pristipomoides auricilLa. A single specimen of P. auricilLa (12.1 mm) has been identified above; it has strong internal structure in the dorsal-fin spines, and very low numbers of spines on the outer border of the preopercle. None of the specimens of species S, the largest of which is 10.3 mm, have any internal structure in the dorsal fin spines, so it seems unlikely that species S is P. auricilLa. Also, Pristipomoides species S larvae have many more spines on the outer border of the preopercle (Fig. 48) than does the single P. auricilLa (Fig. 41). With 69-73 lateral-line scales, an identification of A. furca is indicated by Anderson's (1987) counts, but according to Kami (1973), P. auricilla cannot be eliminated by these counts (it should be noted that the larvae identified as P. sieboldii have lateral line counts very similar to those of Pristipomoides species S). Finally, although the length of the ascending process of the premaxilla in the three largest specimens of species S overlaps somewhat with that of Pristipomoides spp., it is on average, shorter than that of Pristipo- moides spp. (Fig. 8), implying an identification of Aphareus. Therefore, although the evidence points to an identification of A. furca, P. auricilla cannot be totally eliminated as a possibility (Table 3). More larvae, especially specimens larger than 10 mm of both types are needed. PRISTIPOMOIDES SPECIES P. The series does not contain larvae large enough to have scales or countable gill-rakers. Identification is based solely on similarity in morphology and pigment to the larvae of the other Pristipomoides and Aphareus species (Table 3). Atlantic Pristipomoides species (there are three Atlantic species, Table I). PRISTIPOMOIDES AQUILONARIS. Fully scaled specimens have 47-51 lateral-line scales. This eliminates P. macrophthalmus. The two largest specimens have 5-6 + 16 gill rakers, apparently eliminating P. freemani (Table I, Fig. 7). Body depth increases with size in this species, and the largest specimen has a body depth of 33% BL, which is greater than the maximum recorded for P. freemani (28.6%) 58 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994 and very close to the minimum recorded for P. aquilonaris (34.5%). Further, P. freemani adults are not known north of Panama (Allen, 1985), and all the material examined here was from north of Cuba (however, see next paragraph). This confirms an identification of P. aquilonaris. PRISTIPOMOIDES FREEMANI? The single specimen is 11.1 mm, and has 52 lateral- line scales. This eliminates P. macrophthalmus. The specimen has 16 gill rakers, which is two to three more than P. aquilonaris of similar size (Fig. 7). This implies that the larva is P. freemani, as that species has more gill rakers than does P. aquilonaris (Table 1). However, it is possible that this is merely a variant of P. aquilonaris, and the identification must remain tentative until more specimens become available. Unidentified Etelinae.-We have five similar larvae (3.8-5.7 mm), apparently of two species, from the western Coral Sea, western Tasman Sea and East China Sea. They have A 111,8,so are not Randallichthys. They are not the monotypic Aprion which has already been identified. Otherwise, nothing certain regarding identification can be said because of their small size and lack of scales. The larvae differ in pigment, development of head spines and fin-spine morphology from the larvae identified as Pristipomoides and Aphareus, so it seems unlikely they are species of either (Table 2). They are most similar to Etelis larvae but differ primarily in having less pigment. However, only three Etelis species are known from the Indo-Pacific, and three types of larvae identified as Etelis are described here, seemingly eliminating this genus as a possibility. Three possibilities arise: there are undescribed species of Etelis or an undescribed Indo-Pacific genus; we have erred in splitting the identified Etelis larvae into three species; or the unidentified larvae are members of Pristipomoides or Aphareus but differ greatly from the larvae of other species of those genera (our ongoing studies reveal that within the apsiline lutjanid genus Paracaesio, similar morphological differences exist among the species). These larvae could represent Pristipomoides argyrogrammicus and/or P. zonatus which have frequently been placed in a separate genus (incorrectly called Tropodinius, see Anderson, 1987). A more complete series is necessary to choose among these possibilities. The two putative species differ from one another in pigment, size at flexion, and possibly in body depth and fin- spine length.

DESCRIPTION OF DEVELOPMENT Development of all the eteline lutjanids is similar, so a general description covering all is given here. Differences among genera and species are covered in the sections on individual taxa. Morphology.-Eteline lutjanid larvae have 24 myomeres, and are compressed and of moderate body depth. The youngest larvae are somewhat more elongate. Body depth changes gradually through development of a steeper profile to the forehead and concomitant deepening of the trunk myomeres to give a slightly hunch-back appearance. The body is deeper at the pelvic base than at the anus. The gut, which is in the process of coiling in the smallest specimens, also deepens, taking on a more triangular shape. At the same time, relative position of the pelvic fin changes probably due to allometric growth. The pelvic fin appears to move from its initial position at about the level of the middle of the pectoral fin forward to about the level of the base of the pectoral fin (=the level of Dspl). These changes are accomplished well before notochord flexion begins. Subsequently, many species again become more elongate, and the pelvic fin in all species gradually moves LEIS AND LEE: ETELlNAE LARVAL DEVELOPMENT S9 posteriorly relative to pectoral and dorsal fin to end up at the level of Dsp2-3. There is a small gap between the anus and the origin of the anal fin which disappears shortly after flexion is complete. In the available specimens, the gas bladder is generally large but not conspicuous because of its position deep within the trunk musculature and extensive shielding by pigment. Some specimens have exceptionally expanded gas bladders, undoubtedly due to rapid ascent to the surface in the net. The head is moderately large and compressed. The snout length is about the same as the eye width. The snout is initially pointed, but as the head profile becomes more rounded, the snout becomes less pointed. The mouth is large, somewhat oblique and reaches at least the anterior margin of the eye, and may extend to mid-eye. Small, conical, larval teeth first form anteriorly on the premaxillae at about the time the third spine of the dorsal fin is forming. Similar dentary teeth form soon after. A pair of canine teeth in both upper and lower jaws forms at 8-10 mm. The nasal pit bridges over to form two nostrils at ca. 8-9 mm. Spination on the head is moderately developed. All head spines are smooth, and none of the ridges have serrations (the serrations on the preopercular outer border seen late in the larval stage are formed by a large increase in the number of spines along that border). The first spine to form is the one at the preopercular angle at about 3 mm. The sequence of (and size at) formation of the first spine on other bones is interopercIe (3.5--4.2 mm), supracIeithral (4-5 mm), posttemporal (4-6 mm), subopercle (5-15 mm, Table 4). The spine at the angle of the preopercIe is the largest on the head, reaching 4-11 % BL. It gradually becomes engulfed by the border of the preopercIe. The inner border of the preopercIe has as many as 4 small spines. These form from about 3 mm and persist until about 18-22 mm. The outer border of the preopercIe is armed with spines up to the size of the pupil. The first of these forms at about 3 mm, the spines gradually increase in number, but their growth does not keep pace with that of the fish, and they eventually form a serrate border on the preopercIe well before settlement. On the upper limb of the outer border of the preopercIe, the number of spines remains at one only until shortly after flexion after which spines are added at regular intervals. The interopercular spination forms initially as a small spine just dorsal to and approximately parallel to the spine at the preopercular angle. This spine is the second longest on the head. Additional small spines are added to the border of the interopercIe until it develops a serrate edge. The small spines on the subopercIe form somewhat later and this bone too eventually becomes bor- dered by serrations. A single opercular spine appears at 3.4--4.6 mm and is joined by a smaller second spine sometime after 16-21 mm. All the spination of the bones of the opercular series is retained by adult as in the reduced form of serrations with two exceptions: the spines on the inner border of the preopercIe disappear, and the two opercular spines are retained as relatively large spines which support the operculum. A small spine forms on the dorsal postcIeithrum just dorsal to the pectoral base at 4.5-5.7 mm (late flexion to early postflexion stage). It persists until at least 40 mm. The supracIeithrum develops its first spine shortly before flexion and adds small spines until obscured by scales at about 50 mm. A small posttemporal spine appears during notochord flexion, and up to five spines are added along the dorsal ridge until the spines disappear at about 30 mm. The ventral ridge of the posttemporal is initially low and spineless, but a small, single spine forms early in the postflexion stage. A low, smooth, spineless, supraocular ridge develops at about the time of notochord flexion and disappears late in the pelagic period. A low, smooth frontal ridge dorsal and slightly anterior to the supraocular ridge 60 BULLETIN OF MARINE SCIENCE. VOL. 55. NO. I. 1994

Table 4. Development of spines on the subopercle in larvae of eteline lutjanids. Given is the size of the smallest specimen to have a spine on the subopercle; unless noted otherwise, all larger larvae have spination on the subopercle. Values in parentheses are the size of the next smallest available specimen. Three taxa do not have a spine in the largest specimen, but the largest specimen is <7 mm: Pristipomoides sp. P, Unidentified Etelinae I and Unidentified Etelinae 2.

Species Size Remarks

Ete/is sp. D 4.9 (4.6) 4.9 mm is largest specimen P. aquilonaris 5.7 (5.5) some to 10 mm lack spine P. multidensltypus 6.9 (6.7) no spine in a 7.5 mm larva P. filamentosus 7.0 (6.2) some to 10 mm lack spine Ete/is sp. A 7.6 (7.4) some to 8.6 mm lack spine Ete/is sp. C 8.3 (7.7) 8.3 mm is largest specimen Pristipomoides sp. S 8.6 (8.6) some to 10 mm lack spine P. auricilla >12.1 only specimen lacks a spine Aprion virescens 12.3 (11.4) P. sieboldii 14.0 (9.6) Aphareus rutilans 28.8 (15.0)

forms at about 7.5 mm. A low, smooth pterotic ridge forms at ca. 6 mm approximately in line with the supraocular ridge. The maxilla develops at its anterior end a structure which could be interpreted as a spine, but which is regarded here as an acute angle. None of the head spines are serrate at any time. Many of the spiny and smooth ridges on the head develop into the walls of the sensory canals. This includes the preopercle, supraocular, frontal, pterotic and portions of the posttemporal. It is possible that these low ridges and the small spines that arm some of them function to direct flow in relation to the developing neuromasts. The spines of the fins and their development are among the most distinctive characteristics of eteline lutjanid larvae. The first fin element to form is the second spine of the dorsal fin. At about the same time, or shortly thereafter, the pelvic spine forms. Most of the dorsal spines and pelvic rays form before the soft rays of the dorsal fin, the elements of the anal fin and the primary caudal rays begin to form. However, the latter are all present by the time notochord flexion is complete. Flexion begins at about 3.5-4.4 mm and is complete at about 4.3-5.5 mm. With some variation in the timing of formation of first spine of the dorsal fin, the dorsal spines form from anterior to posterior. The ultimate spine of the dorsal fin forms initially as a soft ray, but remains so over only a very short range of BL, and transforms into a spine before flexion is complete or very shortly thereafter. The third anal-fin spine forms initially as a soft ray early in the flexion stage and transforms into a spine during flexion or very shortly thereafter. The pectoral-fin rays are the last to form, and full complements of rays are reached about the same time the scales are formed. The second and, in some taxa, the third dorsal spines, and the pelvic spine and first one or two pelvic rays become very elongate (20-50% BL, see species accounts for details). In most species, the second dorsal spine is initially longer than the pelvic spine. Maximum relative spine length is reached shortly after flexion is complete. At least the first and often the second pelvic rays are longer than the spine. Anal spines are not particularly elongate. All spines are chevron-shaped in cross-section, and the concave surface contains tissue that is often pigmented. A ridge develops basally from the concave surface of the chevron of the pelvic-fin spine resulting in a rough "W" shape in cross-section of the trailing surfaces. The timing and extent of this central ridge vary with taxon, but it never forms before 7 mm, and is present in all taxa LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 61 by 15 mm. This central ridge extends distally with growth and may eventually extend along most of the length of the spine. Fin-spine ornamentation varies among species and developmental stage but serrations, if any, are confined to the trailing arms of the chevron (not the central ridge). The leading edge of the spines is smoothly rounded, without ridges. Scales form relatively early. Depending on species, incipient scales are present from about 6.5-8.0 mm, and full complements of scales from 7.6-9.3 mm. Pigment is generally light. A pigment cap is usually present dorsally on the gas bladder and gut. In most taxa melanophores are present dorsally on the midbrain before 3 mm (however, see Unidentified Etelines). These spread to cover the midbrain. An embedded melanophore is present dorsally over the hindbrain, but can no longer be seen before flexion commences. There is initially one or more melanophores along the ventral midline of the tail, but the subsequent fate of these melanophores varies among taxa. Other pigment varies among taxa. Me- lanophores may be present at certain positions such as cleithral symphysis and urostyle only over species-specific, limited size ranges.

SPECIES ACCOUNTS OF DEVELOPMENT Genus Aphareus Cuvier, 1830 (two species, Indo-Pacific) Diagnosis (note that this is based only on A. rutilans).-MoRPHOLOGY-fin spines smooth externally, developing a "frothy" appearance internally (Fig. 21); scales forming late (after 8 mm); no scales on maxilla; ascending process of premaxilla short (Fig. 8); subopercular spines forming very late (Table 4). PIGMENT-single enlarged melanophore becoming a cluster of melanophores at posterior end of anal-fin base, from approximately 3.5 mm, similar pigment at posterior end of dorsal-fin base; pigment internally on urostyle from 5.7-12 mm; pigment present on urostyle only ventrally; no pigment on forebrain until after 8 mm; no pigment on spine of P2; only pigment on P2 distally on two longest rays and their membranes, but on rays 3 and 4 from 7 mm; dorsal-fin pigment limited to distal portions of membranes anteriorly between longest spines; no series of melanophores on dorsal-fin spines, pigment present only distally on Dsp2 and 3, usually an extension of pigment on membrane; no lower jaw pigment. Remarks.- This description is based solely on larvae of A. rutilans. It is possible the taxon called Pristipomoides species S, below, is actually Aphareus furca. Whether it is or not, it is obvious that development of Aphareus is extraordinarily similar to that of Pristipomoides. This is discussed further under Pristipomoides, below. These two genera are easily distinguished from Aprion and Etelis, but are very difficult to distinguish from each other (Table 2).

Aphareus rutilans Cuvier, 1830.-34 specimens, 3.0-43.0 mm Figures 1-8, Table 5

Material examined.-Hawaii, AMS 1.24998-008, 1.25362-022; NMFSLJ T. Clarke 70.9.31: East China Sea, AMS 1.32084-002, 1.32088-001; MUFS 9500, 9501: Philippine Sea, AMS 1.31538-001: western Pacific Ocean, AMS 1.31814-002: Pacific off the north coast of Irian Jaya, ZMUC DANA 3768, DANA 3768-XVII: western Coral Sea, MCZ GRH 1046: western Tasman Sea, AMS 1.26112-003, 1.26112- 004, 1.26122-001, 1.26124-002, 1.26131-009, 1.26140-002, 1.26141-002, 1.26153-006, 1.26]56-002: eastern Indian Ocean west of Sumatra, AMS 1.32513-001, 1.32514-001; ZMUC DANA 3875-III: western Indian Ocean in the Mozambique Channel, AMS 1.33251-002; ZMUC DANA 3949- V, DANA 3962- IV: western Indian Ocean, AMS 1.33252-003: western Indian Ocean off South Africa, RUSI 40876: west Indian Ocean between Comoro Is]ands and Madagascar, AMS 1.33053-005: No data, AMS 1.32515-001 (DANA, unknown station). 62 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

Table 5. Selected morphometric and meristic characters of Aphareus rutilans larvae. Measurements in mm. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus.

Orbit Body Body Body Preanal Predorsal Head diarn- Snout depth depth Dorsal Anal Pectoral LL 10 Sb length length length length ctef length (P) (A) fin fin fin scales spines spines Preftexion 3.04 1.58 1.12 1.08 0.40 0.34 0.88 0.74 IV 0 0 0 0 0 3.29 1.67 1.15 1.05 0.41 0.28 0.85 0.65 IV 0 0 0 0 0 3.46 1.82 1.24 1.13 0.42 0.34 0.82 0.64 II 0 0 0 0 0 3.50 1.75 1.30 1.10 d d 0.90 0.75 III 0 0 0 0 0 3.51 1.81 1.16 1.06 0.38 0.33 0.94 0.75 IV 0 0 0 0 0 3.63 1.75 1.18 1.15 0.42 0.34 1.00 0.75 III a 0 0 0 0 3.76 1.88 1.28 1.38 0.48 0.44 1.12 0.80 VI a 0 0 I 0 3.84 1.64 1.27 1.05 0.43 0.28 0.81 0.71 IV a 0 0 0 0 3.88 2.06 1.47 1.19 0.47 0.41 1.00 0.81 IV a 0 0 0 0 3.96 2.00 1.44 1.44 0.54 0.40 1.16 0.94 VI i 0 0 0 0 4.08 2.12 1.48 1.52 0.52 0.42 1.22 0.86 VI a a a a 4.19 2.09 1.44 1.41 0.53 0.41 1.22 0.94 VI 0 0 I 0 4.19 2.19 1.58 1.47 0.53 0.41 1.25 1.00 VI 0 I 0 Flexion 4.44 2.19 1.44 1.51 0.53 0.42 1.31 1.00 VI 0 0 4.48 2.32 1.52 1.56 0.60 0.48 1.36 1.00 VII 0 0 4.58 2.39 1.55 1.44 0.53 0.48 1.34 0.94 VIII 0 0 4.96 2.56 1.68 1.64 0.64 0.48 1.44 1.12 IX,12 8 0 0 5.20 2.58 1.83 1.64 0.65 0.43 1.33 0.93 VII i 0 a 5.42 2.93 1.95 1.89 0.65 0.56 1.74 1.33 X,12 III,8 3 0 a Postflexion 5.44 2.88 1.96 1.88 0.66 0.56 1.64 1.24 IX,12 I1I,8 I 0 1 0 5.50 3.28 2.46 2.23 0.87 d 1.74 1.46 X,II III,8 8 0 1 0 5.68 3.20 2.32 2.24 0.76 0.68 1.84 1.44 X,II III,8 3 0 1 0 5.95 3.51 2.24 2.25 0.87 0.56 1.94 1.46 X, 11 III,8 7 0 1 0 6.16 3.60 2.36 2.40 0.84 0.68 1.96 1.56 X,11 III,8 d 0 1 0 6.96 4.24 2.88 2.68 0.96 0.68 2.20 1.52 X,11 III,8 9 0 2 0 7.70 4.24 2.80 2.76 0.92 0.76 2.32 1.80 X,II III,8 12 0 2 0 8.00 4.48 3.04 2.88 1.04 0.76 2.60 2.12 X,II III,8 12 i 2 0 8.07 4.61 3.14 3.23 1.03 0.90 2.58 2.06 X,II III,8 -12 0 3 0 8.40 4.75 2.88 2.95 1.04 0.72 2.68 2.16 X,ll III,8 12 2 0 8.56 4.99 2.96 3.04 1.04 0.88 2.56 1.92 X,II III,8 16 3 0 8.60 5.08 3.08 3.35 1.08 1.02 2.55 1.88 X,II III,8 14 71 2 0 8.80 4.88 3.28 3.28 1.12 1.08 2.64 2.08 X,II Ill,8 14 5 0 9.12 4.99 3.44 3.20 l.l8 0.90 2.92 2.44 X,II III,8 12 -73 2 0 10.38 5.38 3.46 3.68 l.l6 0.96 3.03 2.45 X,II III,8 14 -70 3 0 12.14 6.72 4.26 4.23 1.58 1.49 3.65 2.99 X,II III,8 16 71 4 0 14.99 8.07 5.38 5.00 1.67 1.28 4.23 3.59 X,II I1I,8 16 d 4.5 0 28.75 15.42 9.18 9.51 2.99 2.82 7.88 7.14 X, 11 III,8 16 74 5 7 42.95 24.10 14.60 13.94 4.15 4.32 10.66 9.84 X,l1 Ill, 8 16 70 5 7

Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-fin spines long, especially

P2sp (Fig. 4); Dsp2 > P2sp between approximately 3.5 and 5.5 mm, P2 longer thereafter; frothy internal structure in fin spines after approximately 9 mm, initially weak, becoming strong between 15 and 29 mm; number of spines on preopercular outer border low (Fig. 5); lateral-line scales 70-73 (after ca. 8.5 mm); gill rakers very numerous (Fig. 6); ascending process of the premaxilla short (Fig. 8). PIGMENT-forebrain melanophores after approximately 8 mm; no melanophores on lower jaw until at least 15 mm; melanophore present at or just anterior to cleithral symphysis until approximately 7 mm, but may persist until approximately 9 mm; no pigment on base of PI; pigment on rays 3-5 of pelvic fin after LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 63

Figure 1. Larvae of Aphareus rutilans from the western Tasman Sea. Scale bars = 1 mm. A. 3.6 mm (AMS 1.26112-004). B. 4.2 mm (AMS 1.26112-004). C. 4.6 mm (AMS 1.26122-001), note that there are two melanophores at the cleithral symphysis, an unusual condition for this species.

7 mm; internal urostyle pigment from 5.7 mm until approximately ]2 mm, pigment ventral to urostyle; melanophores at tips of caudal rays after approximately 12 mm; no pigment along dorsal-fin base anterior to posterior cluster until approximately 15 mm. Description.-As development of Aphareus rutilans differs only in minor detail from that of Pristipomoides spp., refer to the description of the latter (below) and the diagnosis of Aphareus. above. 64 BULLETIN OF MARINE SCIENCE. VOL. 55. NO. I. 1994

.: ". ":':: -.":" .

. . "." .

Figure 2. Larvae of Aphareus rutilans. Scale bar = 1 mm. A. 6.0 mm from the East China Sea (AMS 1.32084-002). B. 8.4 mm from the Indian Ocean west of Sumatra (AMS 1.32513-001). Incipient scales are present over most of the body, but are shown in only two areas. C. 10.4 mm from the w;:stern Coral Sea (MCZ GRHI046). Scales cover most of the body, but are shown in only one area. Note that supracleithral and posttemporal spines are damaged. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 65

Figure 3. Larvae of Aphareus rutilans. Both specimens have internal structure in the fin spines which is not illustrated. Note pigment at tips of caudal fin. Scale bar = I mm. A. 12.1 mm from the Pacific Ocean off the north coast of Irian Jaya (ZMUC DANA 3768 mw500). Gas bladder is grossly over- inflated, and position of gas bladder pigment is an artefact. Scales shown in only a small area. Spec- imen is badly faded, and some pigment is undoubtedly missing. B. 28.8 mm from an unknown Indo- Pacific DANA station (AMS 1.32515-001). Only scales along the lateral line shown. Myomeres not shown because they are obscured by scales.

Remarks.-As with most eteline species, the series is reasonably complete up to 9 mm, but very patchy thereafter. Some variability in pigment is apparent, but this is very limited. One specimen has a single, small melanophore on the dorsal surface of the urostyle. Two specimens have two melanophores at the cleithral symphysis rather than the usual one. Before scales form, larvae of A. rutilans are likely to be confused with the minority of P. filamentosus larvae that possess a melanophore at the cleithral symphysis. These two taxa can be distinguished because A. rutilans have non- terminal pigment on the pelvic fin after approximately 7 mm and lack internal structure in the fin spines until after the scales form (Table 3). Larvae of P. sieboldii are very similar to A. rutilans, particularly when in the preflexion stage. The two can be distinguished by the lower jaw melanophore, which A. rutilans lacks, and the relative location of the posterior melanophores on the dorsal and anal fins: in A. rutilans, they are generally directly one above the other, whereas 66 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

...J 40 *~ • • Q) • c: ••• • '0. 30 , • en 0 .. u 00 a~ •. .S; 0 • Qi 20 0 D- o 0 • 0 Aphareus rutilans 15 0 0 .c 10 Preflexion 0 i5l c: 0 Flexion •. Q) Postflexion _ ...J 0 ~ ...J 40 ~ Q) 0 c: 30 OJ •••• • '0. .. .. • • en 0 • • (;j ~ 20 0 0 0 C 0 0 0 "C 0 c: 0 (\j 10 15 .c i5l c: 3 4 5 6 7 8 9 10 11 12 13 14 15 29 43 Q) ...J Body Length (mm)

Figure 4. Larvae of Aphareus rutilans. Note scale breaks. Top. Length of pelvic-fin spine (as percent of body length) in relation to body length. Bottom. Length of dorsal-fin spine 2 (as percent of body length) in relation to body length.

12 • • 10

III Q) c: '0. 8 • •• • en •.. • • III • • '3 6 • u•.. Q) a. .... 0 • •..Q) 4 • • 11. .. Aphareus rutilans .. Preflexion • 2 000_ Flexion a 0(1) mo . POSlllexion. • 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Body Length (mm)

Figure 5. Larvae of Aphareus rutilans. Development of spines on the preopercular outer border (does not include spine at angle). Values for 28.8 mm and 43 mm specimens of 21 and 30 spines, respectively, are not plotted. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 67

30.------~------_:l

26 • A. ruillans • .•• P.sieboldli • P. auricUla 24 o P. species S

rn ~~ 20 • I'll a:: .. 18 • .. .. a .. .. • 16 • .. • 00 • .,0 -0 14 0" '" ..0 .. 12

10 . , 8 10 12 14 16 18 20 22 24 26 28 30 32 34 Body Length (mm)

14 • • • tot rn 12 v" [5'g'D v •• Q)•.. ~ v I'll a:: 10 vvv v v a 8 v P. typus/muhlden9 • P. 'IJamentoaus D P. fOamento8US? 6

8 10 12 14 16 18 20 22 24 26 28 30 32 34 Body Length (mm)

Figure 6. Relation of number of gill rakers on the lower limb of the first arch (not including the one at the angle) to body length in larvae of Aphareus and Indo-Pacific Pristipomoides. Top--Species with high numbers of gill rakers (note scale break); not plotted are a 42 mm A. rutilans with 31 rakers, and six P. sieboldii of 37-54 mm with 18-20 rakers. Bottom-Species with low numbers of gill rakers; not plotted are two P. filamentosus of 36.3-36.7 mm with 13 and 14 rakers, respectively, and a 46.7 mm P. typuslmultidens with 11 rakers. 68 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. 1,1994

17 •...t/) 16 • • Q) ~ as 15 • a: 14 • ••• • • • G 13 ... - • Prislipomoides aquilonaris 12 • " Pristipomoides freemani? 11 , , , , , , , 6 8 10 12 14 16 18 20 22 42

Body Length (mm)

1,0 .. .. ""' v .sE 0.9 v v .. ~ )( 0.8 lIII • E v CD "- .,v Q. 0.7 v '0 v ...... If) v If) vv CD 0.6 • 0 0 VA "- .. Q. :l' A0" 0 0• • Cl 0,5 c:: "lD '6 '\10 c: •• • A, rulilans CD 0.4 0 • .• P. sieboldii If)

, 0.0 , 8 10 12 14 16 18 20 22 24 26 28 30 32 Body Length (mm)

Figure 7 (upper). Relation of number of gill rakers on the lower limb of the first arch (not including the one at the angle) to body length in larvae of two Atlantic Pristipomoides species (note scale break). Figure 8 (lower). Relation of the length of the ascending process of the premaxilla to body length in larvae of Aphareus and some Indo-Pacific Pristipomoides species. Some points represent more than one specimen, and values for specimens larger than 32 mm are not plotted, LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 69 in P. sieboldii the dorsal is posterior to the anal. Larger larvae can be distinguished by gill-raker counts (Fig. 6) and pigment on Pz' Aphareus furea (Lacepede 1801).-The larvae of this species may be those described below under the identification Pristipomoides species S.

Genus Aprion Valenciennes 1830 (monotypic, Indo-Pacific) Aprion vireseens Valenciennes 1830 65 specimens, 2.7-17.9 mm Figures 9-11, Table 6

Material Examined.-Hawaii, AMS 1.23566-017, 1.23570-056, 1.23599-008, 1.23599-009, 1.23603- 011. 1.23818-003, 1.23818-005, 1.23818-006, 1.24998-003, 1.24998-004, 1.24998-005, 1.24998-006, 1.24998-012,1.25000-029,1.25000-030,1.25000-033, 1.25000-034, 1.25000-036, 1.25000-037, 1.25362- 005, 1.25362-006, 1.25650-001; NMFSLJ T. Clarke 71.9.3; USNM 322279, 322280, 322281: East China Sea, AMS 1.32089-001; NSMT-PL 23, NSMT-PL 24, NSMT-PL 25, NSMT-PL 26: western Coral Sea, AMS 1.24883-002, 1.30541-001, 1.30556-002, 1.30557-002, 1.30609-007: western Tasman Sea, AMS 1.26062-002, 1.26112-006, 1.26119-003, 1.26129-001, 1.26130-001, 1.26131-001, 1.26135- 001,1.26140-003,1.31383-001,1.31390-001: eastern Indian Ocean west of Sumatra, ZMUC DANA 3875-II, DANA 3875-III: western Indian Ocean off South Africa, RUSI 40877: western Indian Ocean north of Seychelles, AMS 1.33153-001. Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-finspines with trailing edge serrations, but without internal structure; dorsal-fin spines long (Dsp2 to 50% BL shortly after flexion, Fig. 11); relatively large at flexion (flexion-stage larvae 4.3- 6.2 mm, Table 6); spine at preopercular angle long (6--11% BL); subopercular spines form late (Table 4); PI 17-18; 2 supraneurals; approximately 50 lateral- line scales (after 7.5 mm); maxilla unsealed; scales form early (before 7.5 mm); dorsal fin not notched. PIGMENT-semi-internal melanophores along periphery of lower jaw (until at least 7.2 mm); no dorsal pigment on trunk or tail; initially 3-5 ventral melanophores on tail, none enlarged, these disappearing by completion of flexion; no pigment on forebrain or operculum; pigment on fins limited to Dsp2 and Pz ray 1; urostyle pigment after 5 mm; cleithral symphysis pigment present between 3.2 and 7.5 mm, but only in about 18% of specimens. Deseription.-Body relatively slender initially, becoming deeper, and reaching maximum depth of about 35% BL after flexion. Long snout relatively pointed, more so in smaller larvae. Profile initially concave, becoming flat and shallow (not steep). Mouth large, but not reaching past anterior edge of pupil. Head spines moderately long, except PAsp which is large (reaches MRL-ll % BL-shortly after flexion). PUI 1; PLI 2-3, PLI 2 after 4 mm; PUO 6, PUO 2 after 5.5 mm, PUO 3 after approximately 7.5 mm; PLO 6, PLO 1 after 3.7 mm, PLO 2 after approximately 4.5 mm, PLO 3 after approximately 6 mm. Sb 2, Sb 2 after 12 mm. 10 4, 10 1 after approximately 4 mm, 10 2 after approximately 9 mm, 10 3 after approximately 12 mm.Op 1, after approximately 3.5 mm. Scl 3, Sci 1 after 3.9 mm, Sci 2 after late preflex, Sci 3 after 6--7 mm; Pt 3, Pt 1 after 4-5 mm, Pt 2 after 5-8 mm. F ridge after 7.5 mm. FS after 4.1-4.4 mm. Pe bump after 5 mm, ridge after 6.5 mm. All spines but PLI and PUI present in largest specimen. PLI and Pill present in 12.3 mm larva. Notochord flexion 4-6 (usually approximately 5.3) mm. All fin spines with fine serrations along their trailing edges from shortly after their formation; these begin to weaken after 10 mm, with those on Aspl and 3 disappearing first between 12 and 17 mm. In largest larva, serrations weakening but present on all spines except Aspl and 3. No obvious internal structure in spines. Dsp2 present in least developed specimen, all dorsal elements present before flexion; posterior most spine initially forms as ray, changing to 70 BULLETIN OF MARINE SCIENCE, VOL. 55. NO. I. 1994

Figure 9. Larvae of Aprion virescens from nearshore plankton tows in the Pacific Ocean off the Hawaiian Island of Oahu. Scale bars = 1 mm. A. Composite of several damaged specimens, 3.2-3.4 mm (AMS 1.25000-037, 1.23818-005, 1.23818-006). B. 4.2 mm (AMS 1.25000-034), note serrations on Dsp2. C. 4.4 mm (AMS 1.23566-017), note serrations on P2SP. D. 5.0 mm (AMS 1.23818-003). LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 71

Figure 10. Larvae of Aprion virescens from off Oahu, A from a nearshore plankton tow, Band C from offshore midwater trawls. Scale bars = 1 rum. A. 6.1 mm (AMS 1.23559-008). B. 7.1 mm (AMS 1.24998-004), tiny incipient scales are not shown. C. 12.3 mm (AMS 1.24998-005), specimen is fulIy scaled, but only a representative patch of scales on the trunk is shown; myomeres are omitted because they are obscured by scales; very weak serrations at bases of Aspl and 3 not shown; central posterior ridge of P2 spine not shown. 72 BULLETIN OF MARINE SCIENCE, VOL. 55, NO.1. 1994

Table 6. Selected morphometric and meristic characters of Aprion virescens larvae. Measurements in mm. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus. None of the larvae have any subopercular spines except the two largest which have] and 2, respectively.

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth P,sp Dorsal Anal toral LL PO 10 length length length length eter length (P) (A) length fin fin fin scales spines spines Preflexion 2.75 1.50 1.05 0.80 0.34 0.25 0.65 0.55 0.10 II 0 0 0 0 0 3.08 1.36 1.00 0.98 0.32 0.26 0.78 0.64 0.26 I 0 0 0 0 0 3.19 1.47 1.06 0.84 0.33 0.]9 0.69 0.50 I 0 0 0 0 0 3.20 1.63 1.16 0.94 0.34 0.31 0.72 0.63 I 0 0 0 0 0 3.28 1.64 1.06 1.02 0.42 0.26 0.86 0.76 0.]2 III 0 0 0 2 0 3.35 1.55 1.15 0.85 0.38 0.]9 0.70 0.56 I 0 0 0 0 0 3.44 1.72 1.17 1.03 0.34 0.28 0.69 0.56 0.05 11 0 0 0 1 0 3.44 1.8] 1.17 0.97 0.36 0.34 0.66 0.59 I 0 0 0 I 0 3.44 1.80 1.16 1.04 0.44 0.36 0.84 0.68 0.30 III 0 0 0 2 0 3.48 1.88 1.24 1.32 0.42 0.48 0.96 0.80 0.58 IV a 0 0 3 0 3.56 1.72 1.22 1.13 d d 0.88 0.70 0.27 ill 0 0 0 I 0 3.63 1.81 1.25 0.97 d d 0.66 0.59 11 0 0 0 d d 3.68 1.88 1.28 1.32 0.48 0.46 0.98 0.80 0.56 IV a 0 0 2 1 3.70 1.75 1.15 1.15 0.40 0.25 0.90 0.80 0.30 IV a 0 0 2 0 3.75 1.60 1.25 1.10 0.42 d 0.90 0.75 0.53 IV a 0 0 2 0 3.85 1.80 1.30 1.15 0.56 0.35 1.05 0.90 0.53 IV a 0 0 2 0 3.90 1.90 1.25 1.00 0.44 0.30 0.80 0.60 0.27 IV 0 0 0 2 0 3.94 1.97 1.25 1.25 0.50 0.38 1.16 0.78 0.64 V a 0 0 2 I 4.05 2.00 1.50 1.35 0.55 0.45 1.20 0.95 0.87 V a 0 0 2 0 4.10 1.90 1.35 1.38 0.53 0.43 1.10 0.78 0.73 V a 0 0 2 1 4.15 1.95 1.25 1.20 0.44 0.35 0.85 0.70 0.39 ill 0 0 0 2 0 4.16 2.13 1.48 1.44 0.47 0.42 1.11 0.94 0.64 V a 0 0 2 0 4.20 1.90 1.30 1.25 0.53 d 1.25 0.85 0.76 VI a 0 0 3 1 4.25 1.80 1.30 1.05 0.42 0.35 1.00 0.70 0.53 IV a 0 0 2 0 4.35 1.90 1.28 1.31 0.47 0.38 1.06 0.75 0.50 IV a 0 0 2 0 4.40 2.00 1.35 1.40 0.45 0.35 1.25 0.95 0.81 VI i 0 0 3 0 4.52 2.36 1.48 1.64 0.58 0.52 1.28 0.90 0.94 Vill i 0 0 3 I 4.52 2.13 1.35 1.32 0.53 0.44 1.25 0.84 0.72 V a 0 0 2 0 4.53 2.31 1.56 1.56 d 0.50 1.56 0.98 0.89 VI a 0 0 2 I 4.60 2.15 1.65 1.60 0.55 0.50 1.25 0.90 1.05 VII i 0 0 4 I 4.97 2.97 1.48 1.28 0.50 0.34 1.06 0.84 0.44 III a 0 0 2 0 5.00 2.32 1.66 1.41 0.50 0.44 1.28 0.94 0.78 VII d 0 0 3 I 5.90 2.71 1.74 1.71 0.56 0.53 1.41 1.00 0.94 VII 0 3 I 4.24 2.16 1.40 1.60 0.56 0.46 1.30 0.96 0.92 VI a 0 0 3 I Flexion 4.28 2.04 1.40 1.52 0.58 0.44 1.34 1.08 1.06 VIII i 0 3 1 4.40 2.05 1.50 1.40 0.60 0.45 1.20 0.85 0.78 VI i 0 0 3 0 4.52 2.36 1.48 1.64 0.58 0.52 1.28 0.90 0.94 Vill a 0 0 3 I 4.64 2.40 1.68 1.76 0.64 0.68 1.42 1.04 1.12 IX, 12 11,9 0 4 I 4.65 2.45 1.70 1.75 0.55 0.60 1.40 1.05 1.10 VII 1,1 0 3 I 4.68 2.24 1.72 1.76 0.56 0.54 1.36 1.24 1.06 IX,12 11,9 0 4 I 4.97 2.45 1.69 1.78 0.61 0.53 1.50 1.19 1.20 IX, 12 11,9 8 0 3 1 5.18 2.84 1.82 1.94 0.69 d 1.60 1.30 1.37 IX,12 I1I,8 7 0 3 I 5.28 2.72 1.80 1.84 0.60 0.56 1.52 1.14 1.32 X,II 11,9 i 0 4 I 5.75 2.90 1.94 2.13 0.74 0.65 1.87 1.55 1.47 X,ll IIl,8 i 0 5 1 5.83 2.71 1.82 1.86 0.65 0.55 1.70 1.20 1.36 lX,12 11,9 8 0 3 I 6.12 3.03 1.87 2.25 0.70 0.77 1.87 1.32 1.27 IX, 12 11,9 0 3 I Postflexion 5.43 2.67 1.94 2.11 0.71 0.68 1.86 1.46 1.47 X,11 III, 8 8 0 3 5.74 2.97 2.03 2.38 0.84 0.71 2.00 1.68 1.72 X, 11 I1I,8 8 0 5 5.81 2.90 2.00 2.15 0.76 0,75 1.88 1.38 1.63 X,II ill,8 4 0 d 6.00 3.35 2.13 2.19 d d 1.87 1.35 1.72 X,II I1I,8 9 0 5 6.19 3.29 2.19 2.45 0.81 0.77 2.13 1.68 1.63 X,11 ill,8 5 0 5 LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 73

Table 6. Continued

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth P,sp Dorsal Anal toral LL PO 10 length length length length eter length (P) (A) length fin fin fin scales spines spines 6.45 3.55 2.39 2.58 0.84 0.77 2.13 1.74 1.91 X,II III,8 6 0 4.5 I 6.77 3.61 2.45 2.71 0.94 0.81 2.19 1.84 d X,ll III,8 7 0 d d 7.03 3.77 2.39 2.64 0.90 0.81 2.45 1.94 1.88 X,11 III,8 9 0 6 I 7.06 3.74 2.42 2.58 0.84 0.77 2.13 1.61 1.88 X,II III,8 13 0 4 I 7.16 3.74 2.45 2.58 0.84 0.77 2.39 1.80 1.88 X,l1 Ill,8 11 0 5 I 7.16 3.87 2.45 2.88 0.94 0.84 2.41 1.94 2.03 X,II III,8 8 0 5 I 7.52 4.24 2.72 2.88 1.00 0.96 2.76 2.04 2.16 X,l1 Ill,8 12 0 7 I 7.61 4.26 2.71 2.84 0.88 0.85 2.55 2.05 2.25 X,l1 Ill,8 14 50 5 I 7.92 4.36 2.88 3.04 0.96 0.96 2.80 2.36 2.34 X,II III,8 12 -48 6 I 8.07 4.52 2.90 3.23 1.10 0.90 2.71 2.26 2.30 X,II Ill,8 15 5.5 I 9.86 5.25 3.59 3.74 1.19 1.13 3.23 2.58 2.41 X,ll III,8 15 i 7 2 11.40 d d d d d 3.50 2.70 2.71 X,II Ill,8 18 d 7 I 12.30 6.97 4.26 4.45 1.48 1.42 4.00 3.42 3.22 X,II III,8 17 50 6.5 3 17.93 10.63 6.28 6.02 2.00 1.94 4.52 4.06 3.61 X,l1 Ill,8 17.5 48 12 4 spine by end of flexion. Dsp2 always longest fin spine, reaching MRL (approximately 50% BL, Fig. II) just after flexion. Dspl forms before Dsp4. P2sp second longest, reaching MRL (approximately 30% BL) just after flexion. P2sp present from approximately 3 mm, all elements present before flexion. P2 ray 1 long, to

200% length P2sp in preflexion larvae. All anal elements present during flexion, last spine transforming from ray before flexion complete. Some PI rays present from late flexion, all present by 11 mm. Incipient scales appearing at approximately 7 mm; full set by 8-10 mm. In available specimens, only one spine is present along the posterior edge of the scale, originating anterior to the edge of the scale and projecting above and over the edge. Pigment.-Initially, on the ventral midline of the tail, 3-5 me1anophores present from posterior edge of anal anlage to caudal anlage. Melanophore at anal-fin base disappearing as flexion begins leaving 1-2 posteriorly on peduncle and one at base of ventral caudal rays. These disappearing near end of flexion. During flexion, 1-2 melanophores appear internally at urostyle, settling on concave (dorsal) side forming distinct spot which remains at least to 12 mm. These possibly melanophores from ventral edge of tail. No dorsal pigment on trunk or tail. Least d~veloped specimen with single internal melanophore dorsally on hindbrain, but this no longer visible by 4.5 mm. Between 3--4 mm a dorsal melanophore forms posteriorly on each midbrain hemisphere. These gradually increase in number, by 10 mm covering most of midbrain. No melanophores on forebrain or on the operculum. Specimens smaller than 7.2 mm with series of melanophores at least partially internally along periphery of lower jaw tip (damage in some larger specimens makes maximum size at which jaw pigment retained unclear). Lower jaw with 4-6 individual melanophores or more usually as u-shaped smear of pigment. A small melanophore at cleithral symphysis in 18% of larvae between 3.2 and 7.5 mm, but in no others (however, only one non-Hawaiian larva >7.7 mm is available, see below). This melanophore has only been found in larvae from East China Sea and east coast of Australia (28% of larvae within this size range from these two regions have melanophore). The two larvae available from western Indian Ocean (3.7 and 3.9 mm) lack cleithral melanophore. Pigment at cleithral symphysis not present in Hawaiian material, and small number of larvae available 74 BULLETIN OF MARINE SCIENCE, VOL, 55, NO, "I, 1994

A V Aprion virescens ~ ...J o Preflexion ~ e Flexion Gl C 'Q. • Posttlexion en 50 • a; • • • • • .. •• • •• • 0'" 40 •••• • 0 o~ ~ • • '0 0 " C 0°00 " C\I 30 000 o e 0 • a 0 00 0 " .c. 0 000 0 C. 20 c Gl 0 0 ...J q, 10 0 0 00 '----'-----'---'------L--'----'-----'---i..----!-- •....f'\.• r--'---'----'-----' 3 4 5 6 7 8 9 10 11 12 13 17 18 19 Body Length (mm)

E. sp A • # E. sp C 0 ..J S E. sp 0 E. carbunculus CDc: 40 "5. E. CDruscans • ·0 • C1) 0 , • E. ocula1u8 'iii 30 -. .-. • III • 0 • • ca . "c: 20 CII '0 .c. ,0 C, c: j ,

",12 CD c: '5. C1) 10 :;lil • • ~ 8 c." • 0 • CD • • • •• 0 If. • • • 6 • ••• •• 0 •ID ••• 0 4 ... 0 - • 0 • 0 2 .- .,0 , 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Body Length (mm) LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 75 from other regions precludes any conclusions about its occurrence elsewhere. Pigment on dorsal fin associated with Dsp2 and located along posterior edge of spine within groove of the chevron, and usually along distal 30-50% of the spine as a series. One to several melanophores involved, and two largest specimens have some melanophores on fin membrane distally near anterior spines. Pigment on pelvic fin associated with first ray either outlining ray distally, or on membrane between ray and spine. The pigment on these two fins varies in position and melanophore number, possibly due to damage-it is often absent. No other fins are pigmented. Remarks.-A good series of larvae of this species up to 8 mm is available, but we could obtain only five specimens larger than 8 mm, and one of these is in very poor condition. There is considerable variation in size at developmental stage in A. virescens larvae. This is particularly evident at around the time of flexion, and seems to be connected with condition of the larvae. Those larvae in the best condition are generally larger at developmental benchmarks than are larvae in poor condition. This is probably due to delay in fixation and resulting shrinkage. Many of the larvae were collected incidentally in long midwater trawl tows made for mesopelagic fishes. Consequently, a considerable period may have elapsed between capture and fixation, and this can result in shrinkage of larvae, particularly before the vertebral column is ossified (Leis and Trnski, 1989). The larvae in the best condition, in contrast, were usually taken in shorter plankton tows made for larval fishes, and these were usually fixed very soon after capture thus minimizing shrinkage. This probably explains the very low ratio of Dsp2 length! BL seen between 4 and 6.2 mm BL in some specimens (Fig. 11), all of which were captured in short larval fish tows. As spines do not shrink, a smaller ratio results if the larvae do not shrink. Assuming that shrinkage is solely responsible for the variation in the ratio leads to the implication that shrinkage of 20-25% in BL is possible for larvae within this size range. More preflexion specimens of A. virescens were available than for any other eteline, including a large number of specimens not utilized in this study. This has made it easier to detect this apparent shrinkage problem in this species, but it almost certainly happens in other taxa as well. Aprion virescens is a very widespread species ranging from southern Africa to Japan, Hawaii and Australia; however, the vast majority of the available larvae were from Hawaii and Australia. With one exception, we detected no evidence of geographic variation. A melanophore on the cleithral symphysis was found only in western Pacific (Australian and East China Sea) specimens, but only in 28% of those between 3.2 and 7.5 mm. There were proportionately many more small larvae in the western Pacific material than in the Hawaiian material, so this apparent geographical difference could be no more than an artefact due to un- dersampling of small larvae in Hawaii. The serrations on the fin spines of A. virescens constitute the only external fin ornamentation within the Etelinae. Serrations on fin spines are common among

Figure 11 (upper). Aprion virescens-Length of dorsal-fin spine 2 (as percent of body length) in relation to body length. Note scale break. Figure 12 (lower). Etelis spp.-Top. Length of dorsal-fin spine 2 (as percent of body length) in relation to body length. Relatively few values are plotted because this spine is broken in most specimens. Bottom. Development of spines on the preopercular outer border (does not include spine at angle). 76 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994 the Lutjaninae and Caesioninae, but are not known in the Apsilinae or Paradicich- thyinae (unpubl. data). However, the serrations in Aprion are much finer than those in lutjanines and caesionines, and never occur on the leading edge of the spine as they do in the other two subfamilies. The apparent difference between Aprion and the other genera in formation of the spines of the scales must be viewed with caution due to the relatively small size of the largest Aprion. The difference in the morphology (raised vs. flat) of the first spines to form is probably real, but it is questionable if the apparent difference in numbers of spines is anything more than an artefact due to the small size of the largest Aprion. The largest available pelagic larva of A. virescens (17.9 mm) is considerably smaller than those of Aphareus, Etelis and Pristipomoides spp. This could be an artefact of small sample size, but the number of available specimens of the other species is not significantly greater (and for most species, less) than the number of A. virescens larvae. This suggests that A. virescens may become bottom associated (or at least enter habitats not accessible to conventional towed nets) at a smaller size than do the other species.

Genus Etelis Cuvier, 1828 (three Indo-Pacific and one Atlantic species) Diagnosis.-MoRPHoLoGY-fin spines unornamented either internally or externally; dorsal spines moderately long (Fig. 12), but apparently fragile and frequently broken; Dsp2 longest spine in specimens <40 mm, followed by P2sp, and Dsp3; relatively small at flexion (flexion-stage larvae 3.7--4.5 mm); spine at preopercular angle short (6-9% BL until 8.6 mrn, approximately 5% BL at 14 mm, decreasing to 2-3% BL after 40 mm); subopercular spines form early (Table 4); PI 15-16; 3 supraneurals; 48 to 54 lateral line scales; scales form early (before 7 mm); maxilla scaled (maxillary scales appear between 22 and 40 mm); dorsal fin deeply notched (only just becoming notched at 40 mm). PIGMENT-single melanophore at tip of lower jaw in one species, otherwise, no lower jaw pigment; no dorsal pigment on trunk or tail; initially 1-2 ventral melanophores on tail, one at base of anal fin enlarged, but absent in larvae >4.0 to 6.5 mm, depending on species; melanophores present on forebrain after 4.5 to 5.5 mm, depending on species; melanophore internally on operculum after 4.5 to 6.0 mm, depending on species; no urostyle pigment; pigment on dorsal fin limited to chevron groove of Dsp2 and 3, spreading onto other spines after 7 mm and onto fin membrane sometime after 8.7 mm; pigment on pelvic fin limited to chevron groove of spine and tip of soft ray I; Dsp and P2SP chevron groove pigment a series of cIosely- spaced, elongate melanophores; melanophore (occasionally two or three) at cIei- thral symphysis until about 8 mm. Description.-Body depth relatively constant, 30-37% BL until 8.6 mrn, thereafter, species-specific. Snout short. Profile initially concave, becoming smoothly rounded but steep by flexion. Mouth large, maxilla initially reaching to anterior edge of eye, but to mid-pupil by flexion. Head spines moderately long. PUI I; PLI 3, PLI 2 after 5 mm, PLI 3 after 6-7 mm; PUO > 20, PUO 2 after 4-5 mm, PUO 3 after 5-6 mm (rate species-specific, Fig. 12); PLO > 14, PLO 2 after 4- 5 mm, PLO 3 after 4-5.5 mm (rate species-specific, Fig. 12). Sb up to 10, Sb 1 after 7-8 mm, Sb 2 >8.6 mm, <13.7 mm. 10 up to 7, 10 I after 3.5--4 mm, 10 2 after approximately 6 mm, 10 3 after 8-8.5 mm. Op 2, Op I after approximately 4 mm, Op 2 after approximately 20 mrn. Scl to 9, ScI 1st after approximately 3.5 mm, ScI 2 after approximately 5 mm; Pt to 4, Pt 1 and 2 after 4-5 mm. FS in smallest specimen. F ridge after 7.6-7.7 mm; Pe ridge begins as weak swelling LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 77 at approximately 5 mm, leading to weak ridge almost confluent with supraorbital ridge from approximately 6 mm. Notochord flexion 3.7-4.5 mm. All fin spines smooth and without obvious internal structure. Spines apparently fragile, Dsp2 broken in majority of specimens. Dsp2 in least developed specimen, all fin elements present during or shortly after flexion. Dsp2 always longest fin spine, relatively little change in relative length (30-37% BL) before 8.7 mm (Fig. 12). Dsp 1 present in smallest specimen. Penultimate spine of dorsal fin shorter than ultimate spine from 40 mm. P2sp second longest fin spine, present in least developed specimen, all P2 elements during or shortly after flexion. P2 ray 1 longer than spine but frequently broken, MRL 3.3-7.0 mm (to 135% P2sp), reducing to less than 120% P2sp by 8 mm. All anal elements present from during or shortly after flexion. PI rays forming from late flexion, all present by 8 mm. Incipient scales forming from approximately 6.5 mm; full set by 7.0 mm. The first spines on the scales forming as flat projections from margin of posterior edge of scale in same plane as the scale. Some later-forming spines form anterior to edge of scale, and project up and over edge. These later-forming spines present from approximately 14 mm. Scales appearing on maxilla between 22 and 40 mm. Pigment.-Ventrally on tail a melanophore initially present one or two myomeres anterior to caudal anlage, and one on fin fold near caudal anlage; these disappear by 4 mm. A larger melanophore appears at posterior base of anal anlage at 3.7 mm, and disappears at 4.1 to 6.4 mm, depending on species. Thereafter, no pigment on ventral midline of tail. No dorsal pigment on trunk or tail; no lateral pigment on tail. An internal melanophore present dorsally on hindbrain in smallest specimen, but this no longer visible during flexion. One to three dorsal melanophores present on midbrain posteriorly in smallest specimen, increasing in number and covering most of midbrain dorsally by about 7 mm. First one or two forebrain melanophores appear on anterior margin ventrally at 4.3 to 5.6 mm (depending on species), increasing in number and gradually spreading dorsally and posteriorly to cover most of forebrain anteriorly and dorsally by about 7 mm. Forebrain and hindbrain pigment clusters separate until at least 22 mm. Remarks.-Among the available specimens of Etelis are three gaps in size: no specimens are available between 8.7 and 13.7 mm, between 15.7 and 21.1 mm, and between 22.3 mm and 44.0 mm. These gaps not only limit the detail of the following descriptions, but they also limit the precision of the identifications. All specimens larger than 13.7 mm are identifiable to species, but none of those smaller than 8.7 mm are identifiable (see below). Therefore, the six descriptions below deal separately with three identified taxa based on larger specimens, and with three unidentified taxa based on smaller larvae. Etelis oculatus and E. carbunculus remain pelagic to at least 50 mm, and E. coruscans remains pelagic until at least 22 mm. Thus, in this respect, the species of Etelis are more similar to Pristipomoides spp. than they are to Aprion virescens, which apparently leaves the pelagic environment before it reaches 20 mm.

Etelis sp. A-38 specimens, 3.3-8.6 mm Figures 12, 13, 14, Table 7 Material Examined.-Hawaii, AMS 1.23566-023, 1.23603-016; USNM 322282, 322283, 322284, 322285, 322286: Sea of Japan, AMS 1.32079-001: East China Sea, AMS 1.31796-001, 1.32036-001, 1.32084-001, 1.32094-001; NMST-PL 16, NMST-PL 17-001: Philippine Sea, AMS 1.32093-001: west Pacific near Palau, AMS 1.31783-001: Ceram Sea, AMS 1.32474-001; ZMUC DANA 3783-1, DANA 3783-III: Celebes Sea, ZMUC DANA 3740-IV: Pacific Ocean north of Irian Jaya, ZMUC DANA 3768, DANA 3775-11: west Coral Sea, AMS 1.26399-001: west Tasman Sea, AMS 1.26125-003: Indian Ocean west of Sumatra, AMS 1.32475-001; ZMUC DANA 3885-1, DANA 3885-III. Figure 13. Larvae of Etelis sp. A. Scale bars = I mm. A. 3.5 mm from the Pacific Ocean off the Hawaiian Island of Oahu (USNM 322285). B. 3.9 mm from off Oahu (USNM 322286). Note damaged areas in middle of dorsal fin, and at end of anal fin. C. 5.1 mm from the western Coral Sea (AMS I.26399-001). Note missing eye; length of broken Dsp3 estimated from specimen of similar size. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 79

Figure 14. Larvae of Etelis sp. A. Scale bars = I mm. A. 7.4 mm from off Oahu (USNM 322282). Scales are present, but not illustrated. B. 8.6 mIDfrom the Japan Sea (AMS 1.32079-001). Note grossly inflated gas bladder, and that the only scales shown are those along the lateral line (some of these are missing).

Diagnosis.-Indo-Pacific distribution. MORPHOLOGY-numerousspmes on the outer preopercular border (Fig. 12). PIGMENT-no melanophore on lower jaw; pigment on forebrain from 4.3 mm; melanophore posteriorly at anal fin base only from 3.7 to 4.2 mm; melanophore on ventral midline of caudal peduncle until 4.0 mm; one or two (the latter in five specimens) melanophores at cleithral symphysis, but this pigment absent in most specimens >7 mm. Remarks.-More specimens of a greater range of sizes are available of E. sp. A than of the other two unidentified Etelis species. Only one species appears to be represented in the series, so sp. A cannot be E. radiosus because some of the 80 BULLETIN OF MARINE SCIENCE. VOL. 55, NO. I, 1994

Table 7. Selected morphometric and meristic characters of Ete/is sp. A larvae. Measurements in mm. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus. Omitted are three specimens in poor condition (ZMUC Dana 3740-IV: 4.4-5.2 mm) used for fin spine and pigment only. Subopercular spines absent in all specimens except the 7.60 and 8.64 mm larvae which have a single spine.

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth P,sp Dorsal Anal loral LL 10 length length length length eter length (P) (A) length fin fin fin scales spines Preflexion 3.28 1.72 1.24 1.12 0.44 0.36 0.96 0.76 0.54 III a 0 0 0 3.48 1.76 1.32 1.24 0.44 0.36 1.06 0.84 0.60 V a 0 0 1 3.52 1.76 1.28 1.14 0.46 0.32 1.04 0.82 0.56 V a 0 0 0 3.64 1.76 1.40 1.32 0.46 0.38 1.20 0.88 0.74 V a 0 0 1 Flexion 3.52 1.72 1.28 1.16 d d 1.08 0.76 0.78 VIII a 0 0 1 3.68 1.80 1.28 1.24 0.44 0.38 1.16 0.92 d VIII a 0 0 0 3.81 1.84 1.31 1.28 d d 1.31 0.91 0.58 V a 0 0 0 3.88 2.00 1.40 1.30 0.55 0.35 1.23 1.00 d d 1,9 0 0 1 4.05 2.15 1.58 1.55 0.56 0.47 1.40 1.09 0.87 VIII, 12 11,9 0 1 Postftexion 4.32 2.44 1.64 1.76 0.58 0.50 1.48 1.28 d X, II III,8 0 I 4.55 2.40 1.65 1.70 0.65 0.45 1.60 1.25 d X,II III,8 0 I 4.76 2.48 1.84 1.82 0.64 0.52 1.60 1.32 1.06 X,ll III,8 3 0 1 4.80 d 1.75 1.70 0.70 0.40 1.60 d 1.10 X,ll III,8 d 0 0 4.84 2.48 1.84 1.72 0.64 0.52 1.68 1.32 d X,11 III,8 0 I 5.10 2.90 1.85 1.85 0.60 0.45 1.75 1.35 1.02 X,II III,8 0 I 5.50 3.24 2.03 2.19 0.81 0.57 1.78 1.30 d X,II III,8 d 0 I 5.74 3.12 2.19 2.13 0.69 0.59 2.03 1.44 1.13 X,II III,8 8 0 2 5.76 3.28 2.08 2.24 0.80 0.50 2.16 1.76 1.52 X,II III,8 ~IO 0 I 5.76 3.12 2.08 2.04 0.72 0.52 1.98 1.68 1.42 X,ll III,8 13 0 I 5.81 3.35 2.19 2.20 0.75 0.63 1.95 1.50 1.25 X,II III,8 13 0 2 5.90 3.44 2.35 2.19 0.80 0.60 2.10 1.70 1.40 X,ll III,8 12 0 I 5.95 3.56 2.35 2.16 0.80 0.60 1.85 1.50 d X,II III,8 13 0 2 6.28 3.40 2.34 2.27 0.81 0.64 2.00 1.54 1.52 X,II III,8 -13 0 I 6.32 3.35 2.32 2.10 0.78 0.50 2.00 1.60 1.51 X,II III,8 13 0 I 6.45 3.65 2.35 2.20 0.81 d 2.08 1.71 1.57 X,II III,8 11 i I 6.71 3.74 2.32 2.19 0.80 0.60 2.10 1.70 1.66 X,II III,8 14 -52 I 6.84 3.87 2.58 2.45 0.90 0.60 2.10 1.75 1.68 X,ll III,8 12 0 1.5 7.00 3.87 2.45 2.58 1.00 0.60 2.35 1.80 1.67 X,II III,8 14 ~50 1.5 7.00 3.81 2.64 d 0.95 0.60 2.25 1.80 1.61 X,II III,8 d 0 2 7.05 3.97 2.59 2.56 0.93 0.70 2.27 1.78 1.61 X,II III,8 16 -50 I 7.20 3.92 2.64 2.72 0.96 0.72 2.56 2.08 1.86 X,II III,8 15 ~54 2 7.35 4.00 2.45 2.71 0.90 0.65 2.60 1.95 1.80 X,II III,8 14 -49 1.5 7.60 4.39 2.84 2.75 1.00 0.63 2.50 2.00 1.65 X,II III,8 -13 -50 I 7.80 4.46 2.84 2.84 1.10 0.65 2.50 1.90 1.61 X,II III,8 d ~50 I 7.87 4.26 2.80 2.84 0.95 0.60 2.55 2.10 1.90 X,II III,8 15 -51 3 8.58 4.84 2.71 2.84 1.05 0.65 2.75 2.25 1.98 X,II III,8 d -50 2.5 8.64 4.88 2.90 3.04 1.08 0.76 2.88 2.44 2.00 X,ll III,8 16.5 -52 2

specimens come from Hawaii where this species does not occur. Therefore, it is either E. coruscans or E. carbunculus, both of which occur in Hawaii. With the possible exception of gill raker counts, none of the characters used to distinguish adult Etelis spp. (Allen, 1985; Anderson, 1987) are useful in specimens of the sizes available of species A, C and D, and based on Pristipomoides adult complements of gill rakers are unlikely to be attained until approximately 20 mm or more. When specimens between 8.6 and t 3 mm become available, it is likely that the number of spines on the preopercular border will be the most useful character in establishing identifications. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 81

Figure 15. Larvae of Etelis sp. C from the Pacific Ocean off the Hawaiian Island of Oahu. Scale bars = 1 mm. A. 4.7 mm (NMFSLJ TA. Clarke 71.9.3). Note that third anal-fin and tenth dorsal-fin elements are transforming from rays to spines. B. 6.4 mm (AMS 1.23603-015).

Etelis sp. C-IO specimens, 4.7-8.3 mm Figures 15, 16, Table 8

Material Examined.-Hawaii, AMS 1.23598-034, 1.23603-015; NMFSLJ T. Clarke 71.9.3; USNM 322287,322288: East China Sea, AMS 1.31794-001, 1.32087-001, 1.32090-001: eastern Indian Ocean west of Sumatra, ZMUC DANA 3875-TI: western Indian Ocean between Comoro Islands and Mada- gascar, AMS 1.33241-002. Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-few spines on the outer preopercular border (Fig. 12). PIGMENT-no melanophore on lower jaw; pigment on forebrain from 5.3 mm; melanophore at posterior base of anal fin in smallest specimen until 6.4 mm (with one exception); melanophore on ventral midline of caudal peduncle absent; one or two (the latter in one specimen) melanophores at cleithral symphysis (absent in largest specimen). Remarks.-The same arguments about identification of Etelis sp. A apply to sp. C. Some of the apparent differences between sp. C and sp. A could be artifacts of small sample size and limited available size range of the former. In particular, it is possible a ventral melanophore on the caudal peduncle is present in specimens of sp. C smaller than 4.7 mm. The 5.3 mm specimen from the western Indian Ocean lacks the melanophore at the anal fin base. Too few specimens are available to evaluate if this discrepancy is due to geographical variation. 82 BULLETIN OF MAR[NE SCIENCE. VOL. 55. NO. I. [994

o . . r' ' :,-' ;.;." ", ", " ", ,

Figure 16. Larvae of Ete/is sp. C from the East China Sea; note that although the larvae are fully- scaled, the only scales shown are those along the lateral line, and that some of these are missing. Scale bars = I mm. A. 7.7 mm CAMS 1.32087-001). B. 8.3 mm CAMS 1.31794-001).

Etelis sp. D-3 specimens, 4.4-4.9 mm Figure 17, Table 9

Material Examined.-East China Sea, AMS 1.31540-001, 1.31804-001: Andaman Sea, ZMUC DANA 3896-1.

Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-intermediate number of spines on the outer preopercular border (Fig. 12). PIGMENT--oneor two melanophores on lower jaw; one or two melanophores on forebrain; no melanophore posteriorly on anal-fin base; melanophore on ventral midline of caudal peduncle present only in smallest (4.4 mm) specimen; two or three melanophores at cleithral symphysis. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 83

Table 8. Selected morphometric and meristic characters of Etelis sp. C larvae. Measurements in mm. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus. Subopercular spines absent in all but the 8.3 mm specimen which has one.

Pre- Body Body Pec- Body Preanal dorsal Head Orbit Snout depth depth P2SP Dorsal Anal toral LL 10 length length length length diameter length (P) (A) length fin fin fin scales spines Flexion 4.67 2.58 1.74 1.69 0.59 0.41 1.34 1.16 d IX,12 Il,9 0 Postflexion 4.50 2.35 1.60 1.60 0.60 0.40 1.50 1.10 d IX,12 11,9 i 0 0 5.16 2.88 1.94 1.75 0.63 0.55 1.60 1.40 1.18 X,II ill,8 i 0 1 5.30 2.92 1.91 1.99 0.66 0.55 1.61 1.23 d X,ll ill,8 i 0 I 5.60 2.96 1.92 1.96 0.76 0.48 2.00 1.80 1.34 X,II IIl,8 7 0 I 5.61 3.03 2.06 2.00 0.69 0.44 1.88 1.78 1.26 X,II IIl,8 6 0 1 6.38 3.48 2.39 2.30 0.85 0.60 2.05 1.70 1.36 X,II ill,8 9 0 I 6.39 3.42 2.23 2.26 0.77 0.58 2.00 1.70 d X,II IIl,8 d 0 I 7.68 4.48 2.80 2.88 0.96 0.75 2.64 2.08 1.92 X,II ill,8 12 56 1 8.32 4.56 2.80 2.64 1.04 0.76 2.88 2.36 2.04 X,II IIl,8 16 53 3

Remarks.-Because only three specimens of less than 5 mm are available, and the largest lacks scales, the identification as Etelis is uncertain, and the description very incomplete. All characteristics of the three larvae are consistent with such an identification, but this cannot be confirmed until specimens with scales become available. The possibility exists that these three specimens are merely variants of species A or B, and not larvae of a separate species. If they represent a separate species, they may be E. radiosus, which does not occur in Hawaii. Etelis sp. D may have early development of the subopercular spination, as the largest specimen (4.86 mm) has a small subopercular spine. In Ete/is species A and C, this spine does not form until approximately 7.5 mm. Three smaller specimens (3.9-4.1 mm, AMS 1.31785-001) that cannot confidently be linked with the Etelis sp. D series have additional ventral pigment on the tail and the anterior edge of the anus, but possess the lower jaw pigment and Ete/is Dsp2 pigment. Material transitional between these and the larger three specimens is required to determine their identification.

Etelis carbunculus Cuvier, 1828-3 specimens, 13.7-51.5 mm Figure 18, Table 10

Material Examined.-Hawaii, AMS 1.25361-001: East China Sea, AMS 1.26309-001: Bismarck Sea AMS I.19753-040.

Table 9. Selected morphometric and meristic characters of Etelis sp. D larvae. Measurements in mm. d, damaged; i, incipient; a, anlage; (P), at P I base; (A), at anus. Subopercular spines absent in all but the 4.86 mm specimen which has a tiny one.

Pre- Body Body Pec- Body Preanal dorsal Head Orbit Snout depth depth P,sp Dorsal Anal toral LL 10 length length length length diameter length (P) (A) length fin fin fin scales spines Flexion 4.44 2.20 1.56 1.48 0.54 0.46 1.36 1.12 d Vill 0 Postflexion 4.60 2.46 1.83 1.58 0.62 0.47 1.49 1.21 d X,II IIl,8 0 4.86 2.59 1.94 1.86 0.75 0.50 1.55 1.20 1.08 X,II IIl,8 a 84 BULLETIN OF MARINE SCIENCE. VOL. 55, NO.1, 1994

Figure 17, Larvae of Etelis sp. D. Scale bars = 1 mm. A. 4.4 mm from the East China Sea (AMS 1.31540-001). B. 4.9 mm from the Andaman Sea (ZMUC DANA 3896-1400mw).

Diagnosis.-Indo-Pacific distribution. MORPHOLOGY-11-12 gill rakers, BD 29- 33% BL, 50-52 lateral-line scales. Remarks.-Identification of this species and E. coruscans assumes that adult gill- raker counts are similar to those in specimens of 13 to 50 mm. Counts of the Atlantic species E. oculatus of similar size (below) are 2 to 3 rakers below adult values. If a similar situation exists for Indo-Pacific species, the assumption and the identifications are valid.

Etelis coruscans Valenciennes, 1862-3 specimens, 15.7-22.3 mm Figure 19, Table 11

Material Examined.-Bismarck Sea, AMS 1.19705-039: Pacific Ocean north of Irian Jaya, ZMUC DANA 3752 III: Solomon Sea, MCZ 61276 GRHIOII. Diagnosis.-Indo-Pacific distribution. MORPHOLOGY-I6-17 gill rakers, BD 26- 28% SL, 51-54 lateral-line scales. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 85

Figure 18. Larvae of Etelis carbunculus. Only some patches of scales are shown, both specimens have a full complement of scales on the body. Scale bars = I mm. A. 13.7 mm from the Bismark Sea CAMS1.19753-040). B. 51.5 mm from the Pacific Ocean off the Island of Hawaii (AMS 1.25361- 001). Scales on maxilla not shown.

Remarks.---See remarks under E. carbunculus.

Etelis oculatus (Valenciennes, 1828)-4 specimens, 14.8-49.0 mm Figure 20, Table 12

Material Examined.-Caribbean Sea, MCZ 82540, MCZ 82541: Sargasso Sea, MCZ 61473, MCZ 62156.

Diagnosis.-Atlantic distribution. MORPHOLOGY-I4-16 gill rakers, BD 30-31 % BL, 47-52 lateral-line scales.

Table 10. Selected morphometric and meristic characters of Etelis carbunculus larvae. Measurements in mm. (P), at PI base; (A), at anus. All specimens have D X, 11; A II1,8; PI 16. 13.5 mm specimen has at least 48 scales, some are missing.

Pre- Body Body Body Preanal dorsal Head Orbit Snout depth depth Dsp2 P,sp LL PO 10 Sb length length length length diameter length (P) (A) length length scales spines spines spines 13.70 7.81 4.48 4.61 1.65 0.97 4.39 3.61 2.94 2.84 -48 11.5 2 2 43.95 26.45 15.75 13.55 4.45 3.50 12.90 8.60 6.97 7.61 50 40 7 7 51.50 28.05 16.70 16.60 5.44 3.59 14.90 13.20 7.94 8.20 51.5 28 6 5 86 BULLETIN OF MAR[NE SCIENCE, VOL. 55, NO. I, 1994

Table t t. Selected morphometric and meristic characters of Etelis coruscans larvae. Measurements in rom. (P), at PI base; (A), at anus. All specimens have D X,II; A III,8; PI 15-16.

Prc- Body Body Body Preanal dorsal Head Orbit Snout depth depth Dsp2 P2SP LL PO [0 Sb length length length length diameter length (P) (A) length length scales spines spines spines 15.70 8.80 5.30 5.20 1.70 1.15 4.45 3.90 3.20 3.20 52 12.5 2.5 2.5 21.14 12.04 7.17 6.73 2.24 1.41 6.02 5.12 3.67 4.00 -54 14 5 5 22.29 12.30 7.94 7.43 2.56 1.54 5.76 5.38 4.19 4.06 51 16 6 5

Remarks.-Unfortunately, no specimens of Ete/is from the Atlantic smaller than approximately 15 mm are available, so it can only be assumed that they are similar to the Indo-Pacific species. This is the case for Pristipomoides spp.

Genus Pristipomoides Bleeker 1852 (eight Indo-Pacific and three Atlantic species) Diagnosis.-MORPHOLOGY-Fin spines smooth externally, but developing a "frothy" internal structure (Fig. 21); scales forming early to late depending on species; no scales on maxilla; subopercular spines forming early to late, depending on species (Table 4). PIGMENT-anenlarged melanophore becoming a cluster of melanophores at anal-fin base posteriorly, and from approximately 3.5 mm, similar pigment at dorsal-fin base posteriorly; pigment internally on urostyle after 5- 8 mm, depending on species (size at disappearance highly species-dependent); in most specimens, pigment present on urostyle and both dorsal and ventral to it; pigment on forebrain after 8 mm (except P. aquilonaris where it appears at approximately 5 mm); no pigment on spines of P2 (except P. freemani? which has a series of melanophores on spine); pigment present on P2 only distally on membranes and two longest soft rays (except in two species where pigment appears on rays 3 and 4, but not until approximately 12 mm); dorsal-fin pigment limited to distal portions of membranes anteriorly between longest spines; no series of melanophores on spines of dorsal fin, pigment present only distally on Dsp2 and 3, usually as extension of pigment on membrane; pigment on lower jaw, if present, external only.

Figure 19. Larva of Etelis coruscans. 15.7 mm from the Pacific Ocean off the north coast of Irian Jaya (New Guinea) (ZMUC DANA 3752-III 300mw). Note that the only scales shown are those along the lateral line. Scale bars = 1 rom. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 87

Table 12. Selected morphometric and meristic characters of Etelis oculatus larvae. Measurements in mm. (P), at PI base; (A), at anus. All specimens have D X,I1; A III,8; PI 16-17.

Pre- Body Body Body Preanal dorsal Head Orbit Snout depth depth Dsp2 P2SP LL PO 10 Sb length length length length diameter length (P) (A) length length scales spines spines spines Postflexion 14.80 8.70 4.90 5.10 1.80 1.20 4.40 4.00 2.90 2.80 -52 15 6 2 15.70 9.40 5.70 5.70 1.90 1.60 4.70 3.90 3.40 3.20 49 17 4 3 44.20 26.45 14.30 15.80 4.70 4.70 13.40 11.30 6.90 7.40 47 35 7 9 48.95 31.15 19.20 17.80 5.60 4.70 15.30 13.10 7.10 8.10 -52 37 10 10

Description.-MoRPHOLOGY-Body relatively slender initially, becoming deeper up until flexion. Subsequent changes in body depth species-specific. Snout moderate, initially pointed, becoming more rounded with growth, final shape species- specific. Profile initially concave, becoming flatter as rostral cartilage forms. Mouth large, reaching to near mid-pupil in most species. Head spines moderately large, except PAsp which is large (reaches MRL of up to 9% BL at 5-6 mm). PUI 1-2; PLI 2-3; PO varies among species (Figs. 25, 31, 36, 40, 44, 48). Sb to 14, varies with species. 10 to 11, varies with species. Op 1 after 3.7-4.6 mm. Scl up to 10, varies with species. Pt up to 6, varies with species. One each Scl and Pt before flexion complete. F after 7-8 mm; FS after late preflexion to early flexion. Pe low bump after 4.5-5.2 mm, ridge after 5.6-6.7 mm. All spines but PLI and PUI present in largest specimens. Notochord flexion 4-6 mm. All fin spines smooth, roundly chevron-shaped in cross-section. Fin spines developing internal structure of irregular arches within wings of spine chevron resulting in frothy appearance when viewed laterally. Size at development of internal structure and its eventual strength varies among species. Internal structure best developed in Dsp2-5 and P2sp. Dsp2 and 3 in least developed specimen, all dorsal elements present before flexion complete; posteriormost spine initially forms as ray, transforming to spine by end of flexion. Dsp2 or P2sp longest fin spine; Dsp2 reaches MRL (30-40% BL, depending on species) at about the end of flexion, and remains at MRL or only very gradually decreases; P2SP reaches MRL (ca. 30% BL) somewhat later (6-7 mm) (Figs. 24, 30, 35, 40, 43, 47). Dspl forming after Dsp4.

Figure 20. Larva of Etelis oculatus. 15.7 mm from the Caribbean Sea (MCZ 82540). Note that the only scales shown are those along the lateral line and that gas bladder is grossly inflated. Scale bars = 1 mm. 88 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

------~-~- -- - -=- ~:;:..,;. -- -~

Figure 21. Development of internal structure in the dorsal fin of Pristipomoides and Aphareus. Scale bars in A, D, and G are 0.5 mm. All other scale bars are I mm. A-C. Late-developing, weak structure. Internal structure absent in A and B, and only weakly developed in C. A is Aphareus rutilans, which at this size has spine structure identical to Pristipomoides sieboldii which is illustrated in Band C. A. 8.1 mm (AMS 1.24998-008). B. 14.0 mm (AMS 1.25362-009). C. 26.2 mm (AMS 1.25347-002). D-F. Moderate structure. Internal structure absent in D, moderately developed in E and strongly developed in F. All are Pristipomoides multidens and/or typus, D. 8.3 mm (CSIRO AS06/82LF Stn. 10(1». E. 13.6 mm (AMS 1.198714-015). F. 18,2 mm (AMS 1.26410-001), G-1. Early-forming, strong structure. Internal structure strongly to very strongly developed in all three specimens. All are Pris- tipomoidesfilamentosus. G. 8.3 mm (CSIRO AS06/82LF Stn. 3(2». H. 14.1 mm (AMS 1.23603-012). I. 29.5 mm (AMS 1.25349-001).

P2sp present in least developed specimens, all elements present before flexion. P2 ray 1 long, to 180% P2sp in preflexion larvae. All anal elements present during flexion, last spine transforms from ray during flexion or very shortly thereafter. PI rays forming after late flexion-early postflexion, all present by time scales present. Incipient scales forming from 6,5-8.8 mm; full set present by 7.75-10.0 mm. First spines on scales form as flat projections from margin of posterior edge of scale in same plane as scale. Some of late-appearing spines form anterior to edge of scale, and project up and over the edge, These late-appearing spines not present before approximately 19 mm. Pigment.-Initially, on ventral midline of tail, are 3-5 melanophores from posterior edge of anal anlage to caudal anlage. Melanophore at anal-fin base enlarges, and eventually becomes internal at about time scales form, At most, one remains near middle of peduncle, and a second on caudal anlage. The former is variable in persistence (apparently both within and among species), and may disappear well before flexion begins, or be retained until well after the scales are formed. The latter tends to be retained and becomes located in middle of the base of ventral caudal rays. Internal melanophores appear on urostyle at species-specific times: their persistence is also species specific. In all species but p, filamentosus LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 89

Table 13. Selected morphometric and meristic characters of Pristipomoides aquilonaris larvae. Mea- surements in rom. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus.

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal toral LL 10 Sb length length length length eter length (P) (A) fin fin fin scales spines spines

Preflexion 3.16 1.56 1.20 1.14 0.34 d 0.80 0.66 III a 0 0 0 0 3.52 1.80 1.36 1.32 0.44 d 0.96 0.78 IV a 0 0 0 0 4.06 1.88 1.31 1.19 0.47 0.31 0.86 0.67 III a 0 0 0 0 4.38 2.03 1.56 1.25 0.44 0.42 1.00 0.72 II a 0 0 0 0 4.48 2.13 1.48 1.35 0.52 0.38 1.19 0.83 V i 0 0 I 0 4.68 2.13 1.45 1.29 0.50 0.38 1.05 0.78 V i 0 0 I 0 Flexion 3.70 2.00 1.60 1.60 0.50 0.50 1.20 0.90 VII a 0 0 ] 0 3.76 2.00 1.50 1.40 0.40 0.40 1.10 1.00 V a 0 0 0 0 4.06 2.39 1.61 1.48 0.55 0.47 1.22 1.00 V i 0 I 0 4.42 2.19 1.55 1.39 0.54 0.41 1.11 0.81 V i 0 I 0 4.71 2.45 1.74 1.55 0.55 0.44 1.09 0.91 VII, I I 1,9 0 0 0 5.16 2.63 1.77 1.68 0.58 0.48 1.32 0.97 VIII, I I 9 0 I 0 Post flexion 5.04 2.70 1.90 1.90 0.62 0.68 1.40 1.20 IX,12 11,9 0 0 0 5.20 3.20 2.30 2.00 0.60 0.50 1.50 1.30 X,II Ill,8 5 0 ] 0 5.36 2.96 2.08 2.00 0.74 0.50 0.90 0.76 X,II Ill,8 0 I 0 5.48 2.90 2.00 1.84 0.67 0.50 1.59 1.09 X,11 Ill,8 i 0 I 0 5.68 3.28 2.24 2.16 0.80 0.60 1.80 1.44 X,II Ill, 8 6 0 ] I 5.90 3.40 2.30 2.20 0.80 0.70 1.90 1.40 X,II Ill,8 0 0 I 0 5.90 3.50 2.50 2.40 0.90 0.80 1.80 1.50 X,II Ill,8 7 0 I I 6.24 3.28 2.32 2.24 0.80 0.56 1.88 1.56 X,II Ill,8 6 0 ] 0 6.26 3.36 2.23 2.13 0.75 0.66 1.81 1.47 X,II 1II,8 -6 0 ] 0 6.32 3.36 2.48 2.32 0.84 0.68 1.96 1.64 X,ll 1II,8 13 0 I 0 6.45 3.55 2.24 2.13 0.80 0.52 2.00 1.61 X,II Ill,8 -5 0 I 0 6.77 3.61 2.32 2.26 0.81 0.65 2.00 1.61 X,II III, 8 7 0 ] 0 6.80 3.68 2.64 2.40 0.92 0.64 2.16 1.84 X,II ill, 8 10 0 2 I 7.04 3.84 2.64 2.56 0.92 0.76 2.32 2.00 X,II ill,8 15 0 2 0 7.28 3.92 2.56 2.64 0.96 0.76 2.36 2.04 X,II Ill,8 16 0 2 I 7.36 4.08 2.72 2.56 1.00 0.72 2.48 1.96 X,II ill,8 13 0 2 0 7.52 4.16 2.80 2.64 1.00 0.68 2.52 2.16 X,I! III, 8 9 0 2 0 7.92 4.16 2.96 2.72 1.12 0.72 2.56 2.04 X,II ill, 8 14 0 2 0 8.00 4.70 3.00 3.10 1.08 0.80 2.20 1.80 X,II ill,8 14 0 2 0 8.08 4.72 3.04 3.12 0.96 0.88 2.68 2.24 X,II ill,8 16 0 2 2 8.45 5.10 3.20 3.10 1.00 0.80 2.50 2.10 X,II ill,8 9 0 3 I 8.96 4.99 3.12 3.04 1.20 0.72 2.76 2.48 X, 10 ill,8 16 0 3 0 9.28 5.15 3.20 3.28 1.20 0.80 2.96 2.56 X,II 1II,8 16 49 3 I 9.36 5.15 3.12 3.28 1.20 0.88 3.00 2.60 X,II 1II,8 16 50 3 I 9.82 5.47 3.60 3.44 1.32 0.88 2.84 2.48 X,II III, 8 16 48 3 ] 10.00 5.70 3.60 3.90 1.28 1.00 3.28 2.80 X,II III, 8 16 47 3 0 10.47 5.80 3.70 3.68 1.36 0.92 3.32 2.88 X,II ill,8 IS 49 4 0 10.79 6.12 3.86 3.92 1.36 1.08 3.36 2.80 X,II ill,8 16 51 2.5 I 10.79 6.10 3.86 3.44 1.28 0.88 3.28 2.72 X,ll ill,8 16 51 3 I 10.95 5.96 3.92 3.68 1.28 1.00 3.36 2.88 X,II ill,8 16 50 4 I 11.50 6.40 4.20 4.00 1.50 1.30 3.60 3.20 X,II ill, 8 15 49 3 1 14.43 7.71 4.90 5.08 1.92 1.40 4.56 4.12 X,II III, 8 14 49 4 3 16.73 9.68 5.58 5.08 2.00 1.60 5.40 4.90 X,II III,S' 15 47 5 5 19.00 10.80 6.45 6.75 2.04 1.75 5.75 4.98 X,II III,8 15 49 5 5 21.65 12.14 7.54 7.22 2.28 1.99 6.39 5.31 X,II III,8 15 -49 5 5 42.36 23.94 15.08 13.28 5.20 4.70 13.50 13.30 X,II I1I,8 IS 51 10 8 90 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

Figure 22. Larvae of Pristipomoides aquilonaris from the Gulf of Mexico (A, B) and the Florida Keys (C). Multiple melanophores at tip of lower jaw not always visible in lateral view. Note melanophore at base of caudal anlage in preflexion larvae and which is homologous with the melanophore at the base of ventral caudal rays in flexion and postflexion larvae. Scale bars = I mm. A. 4.1 mm (NMFSM IS7205 sta 112). B. 4.9 mm (NMFSM IS7320 sta 82). Pigment on midbrain is not sym- metrical; that illustrated is from the right hemisphere. C. 6.2 mm (NMFSM CA8910-6708). LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 91

c Figure 23. Larvae of Pristipomoides aquilonaris from the Gulf of Mexico (A, B) and the Sargasso Sea (C). Multiple melanophores at tip of lower jaw not always visible in lateral view. Scale bars = 1 mm. A. 7.4 mm (NMFSM CA8910-7607). Only some of the incipient scales are shown and internal structure in Dsp 1 and 2 is not shown. B. 10.5 mm (NMFSM CA891O-5207). Specimen is fully scaled, but scales are shown only along lateral line; myomeres of the caudal peduncle could not be discerned; internal structure in fin spines is not shown. C. 19.0 rom (MCZ 82537). Specimen is fully scaled, but scales are shown only along lateral line; internal structure in fin spines is not shown. 92 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994 ~ -1 ~ Pristlpomoides aquilonaris ell .!: 40 0 Preflexion a. iI en Flexion 0 • Postflexion Z 30 iI • ell iI • D- • • • •• a. • •• • •• • 0. iI '020 0 • • • 0 .c c. 00 c: ell 10 Pris1ipomoides freemani? ..I •....• 0. Postflexlon

g30 iI iI • ""iii • • • • • VI • • •••• • • •.. 0 • • 20 1.0 iI j' •• • • • •• 0. 0 iI CI 0 0 • • "0 0 c: N 10 '0 .cc. , , , , c: 4 7 ell 3 5 6 8 9 10 11 12 13 14 15 16 ..I Body Length (mm)

• 18 • • • 14 • "0. ; 12 • • .!: Q. • • •• ~ 10 Ol "S • 0 Q).. 8 • • Q. • 0 .. Pris1ipomoldes freemanl? Q) • 0: 6 .-. Do Postflexion • • • PriBtipomoid88 aquilonaris 4 • • • • o Preflexlon •• e Flexion 2

3 4 5 6 7 8 9 10 11 12 14 15 16 17 Body length (mm)

Figure 24 (upper). Larvae of Pristipomoides aquilonaris and P. freemani? Top. Length of pelvic- fin spine (as percent of body length) in relation to body length. Not plotted are values for 16.7, 19.0, 21.7 and 42.4 rum specimens of 21.3, 20.8, 19.0 and 17%, respectively. Bottom. Length of dorsal-fin spine 2 (as percent of body length) in relation to body length. Not plotted are values for 16.7, 19.0, 21.7 and 42.4 mm specimens of 18.9, 17.6, 16.8 and 13.0%, respectively. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 93

Table 14. Selected morphometric and meristic characters of a Pristipomoides auricilla larva. Mea- surements in mm. d, damaged; (P), at PI base; (A), at anus.

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diarn- Snout depth depth Dorsal Anal toral LL 10 Sb length length length length eter length (P) (A) fin fin fin scales spines spines Postflexion 12.14 5.90 4.10 3.98 1.49 1:49 3.65 3.15 X,1O III, 8 16.5 -70 0

(where they are ventral to urostlye) melanophores are more-or-less centered over urostyle. At 3.5--4.4 mm, a melanophore appears at dorsal-fin base posteriorly approximately opposite the anal-fin base pigment. Melanophores gradually increase in number along bases of rays 9-11 and onto caudal peduncle by time scales form. In most species, melanophores spread anteriorly from this cluster sometime after 10 mm. In some species, melanophores appear at bases of anterior one to three D soft rays forming a cluster well before posterior cluster spreads anteriorly. Least developed specimens have on hindbrain a melanophore dorsally which gradually sinks under anterior dorsal musculature. Least developed larvae have one or two melanophores posteriorly on dorsal surface of each midbrain hemisphere. These gradually increase in number, but remain in posterior half of midbrain until about 8 mm after which they spread anteriorly to cover midbrain. Melanophores form on forebrain at species-specific sizes ranging from 5.2-10.1 mm. One to three melanophores form on internal surface of operculum near the base of opercular spine from late flexion stage onward. Pigment on tip of lower jaw varies with species, but if present, is external consisting of discrete melanophores. Cleithral symphysis pigment species-specific: one or more melanophores may be present at or just anterior to cleithral symphysis, but begin to disappear at about 7 mm, and are absent in all specimens by approximately 10 mm. Pigment on dorsal fin limited to distal portions of membranes, initially between Dsp2, 3 and 4. This may spread posteriorly to all inter-spinous membranes, but remains limited to distal half. The extent of this pigment is species specific. Especially in specimens with torn fins, some of this pigment may appear to be located in chevron groove of spines. This may be an artefact. Presence and extent of pigment on dorsal fin depends very much on condition of specimen. Pigment on pelvic fin limited initially to distal portions of rays 1 and 2 and associated membranes. In some species, pigment may spread to other portions of fin rays and membranes as larvae grow. The single exception is P. freemani? where a series of melanophores is present in the chevron groove of P2sp. In some species, latger larvae have pigmented tips on caudal fin. Neither A nor PI fin pigmented. Remarks.-Individuals of Pristipomoides remain pelagic to considerable size (adults may be semi-pelagic, feeding well off the bottom; W. J. Richards, pers. comm.). During this study, pelagic individuals exceeding 30 mm BL of four species were examined: P. filamentosus (to 36.7 mm), P. aquilonaris (to 42.4 mm), P. multidens and/or typus (to 46.7 mm), P. sieboldii (to 53.8 mm). It is probably not coincidental that these were the four Pristipomoides species for

Figure 25 (lower). Larvae of Pristipomoides aquilonaris and P. freemani? Development of spines on the preopercular outer border (does not include spine at angle). Note scale breaks in both horizontal and vertical axes. Not plotted are values for 19.0, 21.7 and 42.4 mm specimens of 20, 23 and 49 spines, respectively. 94 BULLETIN OF MARINE SCIENCE. VOL. 55. NO. I. 1994

Table 15. Selected morphometric and meristic characters of Pristipomoidesfilamentosus larvae. Mea- surements in mm. * Specimens with melanophore at cleithral symphysis. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus.

Pre- Orbil Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal toral LL 10 Sb length length length length eter length (P) (A) fin fin fin scales spines spines Preflexion 2.93 1.38 1.05 0.65 0.25 0.20 0.65 0.55 a 0 0 0 0 0 3.15 1.65 I.I I 0.85 d d 0.75 0.60 a 0 0 0 0 0 3.25 1.50 1.10 0.95 0.35 0.20 0.85 0.68 II 0 0 0 0 0 3.35 1.70 1.20 0.98 0.35 0.30 0.85 0.70 II 0 0 0 0 0 3.55 1.90 1.40 1.25 0.40 0.35 0.88 0.68 III 0 0 0 0 0 3.64 1.69 1.23 1.03 0.36 0.30 0.72 0.64 II 0 0 0 0 0 3.80 1.70 1.30 1.20 0.40 0.40 1.00 0.60 III 0 0 0 0 0 3.88 1.79 1.46 1.21 0.46 0.35 0.87 0.71 II 0 0 0 0 0 4.00 1.90 1.35 1.05 0.35 d 0.95 0.75 IV 0 0 0 I 0 *4.00 1.98 1.39 1.31 -0.53 0.38 1.22 0.88 IV 0 0 0 0 0 4.00 2.06 1.44 1.22 0.40 0.35 0.84 0.63 II a 0 0 0 0 4.10 2.00 1.50 1.30 0.45 0.45 1.05 0.85 IV a 0 0 0 0 4.40 2.19 1.44 1.38 0.50 d 1.06 0.84 m a 0 0 0 0 4.45 2.00 1.35 1.38 0.40 0.43 1.20 0.95 IV a 0 0 0 0 4.64 2.52 1.61 1.55 0.52 0.45 1.32 1.10 VII 0 0 0 1 0 Flexion *3.54 2.10 1.66 1.79 0.54 0.65 1.20 0.97 V i 0 1 0 *3.94 2.25 1.51 1.61 0.57 0.48 1.31 0.98 VII i i 0 I 0 4.00 2.13 1.52 1.45 0.48 d 1.10 0.97 V 0 0 0 0 0 *4.12 2.34 1.58 1.67 0.58 0.44 1.42 1.06 VIII,2 II 0 I 0 *4.12 2.49 1.85 1.71 0.57 0.46 1.54 1.18 VII 11,9 0 I 0 *4.49 2.71 1.72 1.70 0.61 0.49 1.51 1.12 X,ll 11,9 0 1 0 4.51 2.45 1.71 1.61 0.58 0.39 1.48 1.13 VIII,2 1,7 0 I 0 *4.55 2.84 1.89 1.95 0.63 0.50 1.47 1.13 X,ll 11,8 0 1 0 *4.68 2.44 1.76 1.76 0.60 0.52 1.36 1.08 IX,12 11,9 i 0 I 0 4.71 2.68 1.87 1.87 0.65 0.52 1.55 1.23 IX,12 11,9 0 0 I 0 *4.91 2.71 1.82 1.98 0.57 0.61 1.52 1.10 X,11 III,8 0 I 0 *5.06 3.02 1.91 1.99 0.74 0.50 1.52 1.26 X,11 III,8 0 I 0 *5.10 2.56 1.76 1.76 0.64 0.50 1.30 1.04 X,II III,8 0 I 0 5.29 3.10 2.06 1.97 0.62 0.58 1.74 1.35 X,11 III,d -5 0 I 0 *5.61 2.97 2.00 2.00 0.71 0.58 1.81 1.32 X,d III,8 0 I 0 Postflexion 4.75 2.70 1.75 1.75 0.60 0.50 1.50 1.15 IX,12 11,9 0 0 I 0 5.29 2.84 1.94 1.81 0.71 0.48 1.74 1.42 X,II III,8 0 0 I 0 5.56 2.91 1.91 1.91 0.65 0.50 1.74 1.41 IX,12 III,9 0 0 1 0 *5.84 3.16 2.26 2.26 0.75 0.63 1.75 1.44 X,II III,8 -3 0 I 0 6.14 3.74 2.32 2.40 0.92 0.76 1.96 1.44 X,ll III,8 -6 0 2 0 6.22 3.48 2.26 2.19 0.84 0.58 2.06 1.58 X,ll IlI,8 -8 0 2.5 0 6.97 4.15 2.66 2.82 0.92 0.80 2.04 1.68 X,II IlI,8 0 0 2 I 6.97 3.90 2.57 2.57 0.99 0.91 2.24 1.83 X,II III,8 -9 0 1 0 7.14 4.15 2.82 2.66 0.99 0.83 2.32 1.91 X,ll III,8 -9 0 3 0 7.16 4.00 2.64 2.71 0.90 0.71 2.19 1.77 X,II III,8 II 0 2 1 7.17 4.23 2.75 2.65 0.90 0.81 2.26 1.81 X,II m,8 d 0 2 0 7.64 4.32 3.07 2.99 1.08 0.75 2.64 1.96 X,II III,8 12 0 2 0 *8.08 4.88 2.96 2.88 1.04 0.78 2.76 2.32 X,l1 III,8 13 62 3 0 8.13 4.77 2.90 2.84 1.01 0.65 2.51 2.03 X,II m,8 10 -60 2 0 8.26 4.87 2.95 3.20 1.03 0.84 2.52 2.06 X,II III,8 -10 -60 3 0 8.38 4.91 3.46 3.19 1.08 0.69 2.64 2.14 X,11 m,8 17 -61 3 I 8.45 4.74 3.03 2.97 1.00 0.77 2.64 2.13 X,ll III,8 15 i 3 1 8.58 5.06 3.08 3.16 1.05 0.85 2.67 2.15 X,ll III,8 -14 -62 3.5 0 8.84 4.93 3.20 3.20 1.10 0.84 2.58 2.10 X,II m,8 14 i 3 I 9.20 5.36 3.36 3.28 1.04 0.92 2.92 2.52 X,II III,8 15 64 3 I 9.60 5.19 3.46 3.29 1.10 0.84 3.23 2.71 X,11 III,8 13 d 3 I 10.00 5.81 3.55 3.74 1.23 0.90 3.16 2.58 X,II III,8 15 d 3 0 LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 95

Table 15. Continued

Pre- Orbit Body Body Pee- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal tor•..1 LL 10 Sb length length length length eler length (P) (A) fin fin fin scales spines spines *10.12 5.76 3.59 d 1.16 0.87 3.35 2.58 X,II III,8 d 63 4 I *10.89 5.89 3.78 3.90 1.23 0.90 3.35 2.77 X,11 III, 8 d -63 4 I 11.64 d 4.48 3.90 1.41 1.01 3.65 d X,11 III, 8 16 -59 5 1 11.66 6.41 4.04 4.10 1.35 0.97 3.74 2.90 X,12 III, 8 16 61 4 2 12.30 7.22 4.26 3.61 1.16 1.16 3.98 3.49 X,11 III,8 15 64 5 4 12.46 7.05 4.48 4.48 1.58 1.33 4.32 3.40 X,11 III, 8 15 62 4 4 12.79 d 4.65 4.32 1.58 1.25 3.98 d X,ll TTl, 8 16 60 5 2.5 14.09 8.07 5.12 5.00 1.48 1.35 4.39 3.68 X,11 III, 8 15.5 -63 5 4 16.90 9.48 5.76 5.64 1.79 1.28 5.00 4.48 X,11 TTl, 8 16 -62 4 3 19.0910.12 6.21 6.41 2.05 1.41 6.08 5.38 X, 11 III, 8 15 -58 4 4 26.10 15.15 9.35 9.18 2.95 2.95 7.54 6.89 X,11 1lI,8 16 61 6 9 29.4617.68 10.63 9.54 3.07 2.31 8.45 7.69 X,11 III,8 16 61.5 6 8 29.5017.00 10.90 8.97 2.80 1.92 7.81 7.43 X,II 1lI,8 16 62 6 9 32.30 18.83 11.28 10.50 3.20 2.50 8.84 7.94 X,II III,8 15.5 62 6 6 36.30 21.05 12.55 12.55 3.98 3.65 10.60 9.30 X,II 1lI,8 16 60 7 5 36.7021.15 13.45 13.12 4.07 4.15 11.15 10.00 X,II III,8 16 62 7 8 which the most material was available, and it is likely all species of the genus are pelagic to similar sizes. Larval development in Pristipomoides and Aphareus is essentially identical (Table 2). Particularly noteworthy characters include the cluster of melanophores

Figure 26. Larvae of Pristipomoides filamentosus from nearshore plankton tows in the Pacific Ocean off the Hawaiian island of Oahu. Scale bars = I mm. A. 3.6 mm (AMS 1.23570-055). B. 4.4 mm (AMS 1.25000-045). 96 BULLETIN OF MAR[NE SCIENCE, VOL. 55, NO. I, 1994

Figure 27, Larvae of Pristipomoides filamentosus from offshore midwater trawls in the Pacific Ocean off Oahu. Scale bars = I mm. A. Composite of two specimens: 4.5 mm (head, AMS 1.25000-041) and 4.0 mm (body, AMS 1.25759-002). The latter is obviously somewhat shrunken due to delay in preservation (see Methods) and was probably ca. 4.5 mm before shrinkage. B. 4.9 mm (specimen lost). Anterior-most ray in both dorsal and anal fins transforming to spines.

at the dorsal fin base posteriorly, and the internal structure of the fin spines. This indicates a close relationship between the two genera. The larvae of the species of Pristipomoides are distinguished by minor pigment differences, the lengths of some fin spines, a few meristic characters and the strength and timing of the appearance of the internal structure of the fin spines. Aphareus rutilans larvae are LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 97

Figure 28. Larvae of Pristipomoides filamentosus from offshore midwater trawls. Strong internal structure in all fin spines not illustrated. Scale bars = I mrn. A. 8.3 mrn (CSIRO AS06J82 LF Stn. 3(2)} from the eastern Indian Ocean over the northwest shelf of Australia. Incipient scales cover most of body, but are shown in only a small area. B. 10.0 mm (AMS 1.23598-001) from the Pacific Ocean off Oahu. Lengths of broken spines estimated from a 10. I mm specimen. Note very faint melanophore ventral to the urostyle. Scales cover the entire body, but are illustrated in only a smaJl area. distinguished from Pristipomoides spp. larvae by the same sorts of characters (Table 3). The principal distinguishing pigment character is the presence in A. rutilans of pigment on rays 3-5 of the pelvic fin in larvae >7 mm. However, this pigment pattern occurs also in the larger larvae (>12 mm) of at least two Pris- tipomoides spp. (filamentosus and multidens/typus), so the difference is merely one of timing. Only the length of the ascending process of the premaxilla (long 98 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I. 1994

Figure 29. Larvae of Pristipomoides filamentosus from offshore mid water trawls. Internal structure in all fin spines not illustrated. Scales cover the entire body, but are illustrated in only a small area. Scale bars = 1 mm. A. 16.9 mm (AMS 1.25348-001) from the Pacific Ocean off the island of Hawaii. Length of pelvic rays estimated from 14.1 and 19.1 mm specimens. B. 32.3 mm (AMS 1.24163-008) from the western Tasman Sea. Blank area at base of caudal rays indicates coverage by scales.

Figure 30 (upper). Larvae of Pristipomoides filamentosus. Top. Length of pelvic-fin spine (as percent of body length) in relation to body length. Not plotted are values for eight 16.9-36.7 mm specimens of 26.3-21.3%, respectively. Bottom. Length of dorsal-fin spine 2 (as percent of body length) in relation to body length. Symbols (squares) labeled as Pristipomoidesfilamentosus? represent specimens with a melanophore at the cleithral symphysis. Not plotted are values for eight 16.9-36.7 mm specimens of 23.2-17.6%, respectively. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 99

Ii....I S Ql c: '0. 40 Cf.I 0 .;; & • • • • 30 & &. ., • • • Gi , ••••••• • •• • ell 0. & a i5 20 II a s:. a CD o°q, c: a a ....IQl 10 00 Ii....I S 40 Ql • lI. c: '0. & II lI. • • Cf.I 30 II •••••• • II • • • • • a; 00 • • r/J •• c c 0 0 ';< 5 20 00'0 a ';< c 0 .!! 0 ,! "0 a; ;; a .. 0 c: Q. "u:: 0. (\j a 10 a II • P. flIamento8uS i5 a c & P. filBmentosus? s:. • CD c: Ql 7 8 ....I 3 4 5 6 9 10 11 12 13 14 15 16 Body Length (mm)

16 • 14 •

r/J • Ql 12 • • c: '0. Cf.I •... 10 • • • a:l '3 0 Qj • - Co a • 0 • • Ql •• •... • • 0. 6 • • • c c 0 •• .2 ';( & ~~s II ~.!~ 4 '"'" o.u.o. & . 51 4iIQ8 o e • P. 1Uamentosus & D iii • P. filamento8US? 2 iii III 00

008,)0

3 4 5 6 7 8 9 10 11 12 13 14 15 16 Body Length (mm)

Figure 31 (lower). Larvae of Pristipomoides filamentosus. Development of spines on the preopercular outer border (does not include spine at angle). Symbols (squares) labeled as Pristipomoides filamentosus? represent specimens with a melanophore at the cleithral symphysis. Not plotted are values for eight 16.9-36.7 mm specimens of 16-36 spines, respectively. 100 BULLETIN OF MARINE SCIENCE. VOL. 55. NO. I. 1994

Table 16. Selected morphometric and meristic characters of a Pristipomoides freemani? larva. Mea- surements in rom. (P), at PI base; (A), at anus.

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal toral LL 10 Sb length length length length eter length (P) (A) fin fin fin scales spinesspines 11.10 6.10 3.84 3.52 1.28 0.95 3.36 2.88 X,l1 Ill,8 16 52 2 in Pristipomoides spp., short in A. rutilans), can be regarded as a real difference between the two genera. Identification of Pristipomoides and Aphareus larvae can be difficult because all the species are so similar. Table 3 contains information that is useful in sep- arating larvae of the Indo-Pacific members of these two genera.

Pristipomoides aquilonaris (Goode and Bean, 1896) 48 specimens, 3.2-42.4 mm Figures 22-25, Table 13 Material Examined.-All western Atlantic. Gulf of Mexico, NMFSM 8C7120 sta 34, 8C7120 sta 49, IS7205 sta 64, IS7205 sta 96, IS7205 sta 112, IS7205 sta 114, IS7313 sta 32, IS7320 sta 82, IS7320 sta 98, TIl21 sta 180: Florida Keys, NMFSM CA891O-4207, CA8910-4309, CA891O-5109, CA891O- 5206, CA8901-5207, CA891O-5208, CA891O-5301, CA891O-5806, CA891O-5906, CA891O-6708, CA891O-7304, CA891O-7408, CA891O-7508, CA89 10-7607, CA89 10-7608, CA8910-7708: off SE USA coast (Florida, Georgia, South Carolina), USNM 279422, 279423, 279425, 279477, 279478, 279479, 279480, 279481, 279482, 279483: off Cuba, MCZ 90655: Sargasso Sea, MCZ 82537. Diagnosis.-Western Atlantic distribution. MORPHOLOGy-finspines moderately large (Fig. 24); Dsp2 never longer than P2sp; internal structure present in D and P2 fin spines from approximately 7 mm, becoming moderately strong by approximately 8.5 mm; number of spines on outer border of preopercle high (Fig. 25); subopercular spines form early (Table 4); scales form late (after 9 mm); lateral- line scales 47-51 (after 9 mm); relativelY few gill rakers (Fig. 7). PIGMENT- melanophores present on forebrain from approximately 5 mm; 1-6 melanophores on tip of lower jaw (>90% with 2 or more); a melanophore present on cleithral symphysis until approximately 9 mm; urostyle pigment present from approximately 7 mm; no pigment on base of PI; melanophores present at tips of caudal rays from at least 10.5 mm; no pigment on P2sp; a few melanophores present anteriorly along base of soft dorsal fin in most specimens from approximately 10mm. Remarks.- The series lacks larvae smaller than 3 mm, and although reasonably complete, includes only six individuals larger than 11 mm. We cannot be sure at what size the pigment on the tips of the caudal rays is first present because all specimens between 7 mm and 10 mm have broken caudal-fin rays. This species has, by far, the earliest occurrence of forebrain pigment among Pristipomoides spp.

Pristipomoides auricilla (Jordan, Evermann and Tanaka, 1927) I specimen, 12.1 mm Table 14, Figures 6, 8, 41

Material Examined.-Western Pacific Ocean east of Taiwan, ZMUC DANA 3723 V-50. Diagnosis.-Indo-Pacific distribution. MORPHOLOGY-lengthof fin spines unknown; strong internal structure present in fin spines; number of spines on the outer border of preopercle very low (Fig. 41); subopercular spines form late (Table LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 101

Figure 32. 11.1 mm larva of Pristipomoidesfreemani? (NMFSM CA 8910-5108) from the Western Atlantic Ocean in the vicinity of the Florida Keys. Pigment shown on pelvic spine is that of the right fin; the left fin has only three melanophores. The specimen is fully scaled, but only scales along the lateral line are shown. Internal structure in the fin spines is not illustrated. Scale bar = 1 mm.

4); lateral-line scales approximately 70; moderate numbers of gill rakers (Fig. 6). PIGMENT (due to faded pigment of the only specimen, any notation of absent pigment must be regarded as questionable)-pigment absent on forebrain, lower jaw, urostyle, P2sp, base of PI' and along dorsal-fin base anterior to posterior cluster of melanophores; caudal rays broken. Remarks.- The single available specimen is faded and in relatively poor condition with most of the fin spines broken, so it is not illustrated. It has all the Pristi- pomoideslAphareus characteristics. There is no pigment at either the cleithral or dentary symphyses, or along the base of the soft dorsal fin anteriorly. The lateral- line scale count of 68-72 (the condition of the larva precludes a more precise count), the moderate gill-raker count of 15 (Fig. 6), and long premaxillary ascending process (Fig. 8) eliminate all possibilities except two Pristipomoides species: auricilla and sieboldii. The strong internal structure in the fin spines and very low number of opercular-series spines eliminates sieboldii as a possibility, because sieboldii has no or weak internal structure in the dorsal-fin spines until over 20 mm and has many more opercular-series spines (Fig. 40). This leads to an identification of auricilla. However, the lack of a full series of these larvae and a gap in the sieboldii series between 9 and 14 mm introduces some doubt.

Pristipomoides filamentosus (Valenciennes, 1830) 68 specimens, 2.9-36.7 mm Figures 26-31, Table 15

Material Examined.-Hawaii, AMS 1.23570-053, 1.23570-054, 1.23570-055, 1.23598-026, 1.23603- 012, 1.24998-007, 1.25000-039, 1.25000-040, 1.25000-041, 1.25000-042, 1.25000-045, 1.25348-001, 1.25349-001, 1.25362-007, 1.25362-021, 1.25362-023, 1.25650-002, 1.25759-001, 1.25759-002; NMFSU T. Clarke 71.9.3; USNM 322275, 322276, 322277, 325133: East China Sea, AMS 1.31530- 004,1.31811-001,1.32089-003; ZMUC DANA 3723-IV, DANA 3723-V, DANA 3727-ffi, DANA 3727-IV: South China Sea, AMS 1.26308-001: Celebes Sea, ZMUC DANA 3740-IV: Pacific Ocean north ofIrian Jaya, ZMUC DANA 3768-XVI, DANA 3768-XVII: Bismark Sea, AMS 1.19707-029: western Tasman Sea, AMS 1.17874-021, 1.24163-008, 1.26096-004, 1.26111-012, I. 26124-003, 1.26129-002, 1.26156- 003: Indian Ocean over northwest shelf of Australia, CSIRO AS06/82 LF Stn.3(2): Indian Ocean southwest of Sumatra, ZMUC DANA 3875-11, DANA 3875-ffi, DANA 3885-IV, DANA 3887-IV, DANA 3892-ffi: western Indian Ocean between Comoro Islands and Madgascar, AMS 1.33053-004. 102 BULLETIN OF MARINE SCIENCE, VOL. 55, NO.1, 1994

Figure 33. Larvae of Pristipomoides multidens and/or typus. A from the Gulf of Papua; B, C from the eastern Indian Ocean over the northwest shelf of Australia. Scale bars = I mm. A. 4.1 mm (AMS 1.24552-005). B. 4.8 mm (AMS 1.26412-001). C. 5.7 mm (CSIRO AS06/82LF Stn. 7(3». LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 103

Figure 34. Larvae of Pristipomoides multidens and/or typus. A from the western Tasman Sea; B, C from the eastern Indian Ocean over the northwest shelf of Australia. In A and B incipient scales cover most of the body, but only a few are illustrated. All three illustrated larvae have only 10 gill rakers on the lower limb of the first arch. Scale bars = I mm. A. 7.9 mm (AMS 1.26141-002). B. 8.3 mm (AMS 1.26413-001). C. 18.2 mm (AMS 1.26410-001). Scales cover the body, but are illustrated in only a small area; internal structure in the fin spines is not illustrated. 104 BULLETIN OF MARINE SCIENCE, VOL, 55, NO, I, 1994

:::J al ~ Q) 40 c: '5. en . • • 0 30 . .. •••• ';;: Qg • • .. • Q . •• 0; Q 11. 20 '0 Pristlpomoldes typus/multldens £ Cl c: 10 Flexion Q) ,...,....I Postflexlon ....I al 40 ~ Q) c: '5. 30 en ... • ...... •• g .. , a;

20.------,

16 • 14

Sl 12 c: '5. • • • ~ 10 III "3 o ~ 8 .. c- o Ql•.. •• • 11. 6 ••• • •• • Prlstipomoldes typus/muilidens • 4 Flexion Q •• • • Postflexlon 2 •

3 4 5 6 7 8 9 10 11 12 13 14 15 16 Body Length (mm)

Figure 35 (upper), Larvae of Pristipomoides multidens and/or typus. Top. Length of pelvic fin spine (as percent of body length) in relation to body length. Not plotted are values for five 18.2-46.7 mm specimens of 24.8-19.2%, respectively. Bottom. Length of dorsal fin spine 2 (as percent of body length) in relation to body length. Not plotted are values for five 18.2-46.7 mm specimens of 20.0- 15.2%, respectively. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 105

Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-finspines long (Fig. 30);

Dsp2 > P2sp until approximately 8 mm; internal structure present in fin spines from approximately 4 mm, becoming strong by 6-7 mm; number of spines on outer border of preopercle low (Fig. 31); subopercular spines form early (Table 4); scales form from 8 mm; lateral line scales 60-64 (after ca. 8 mm); relatively few gill rakers (Fig. 6). PIGMENT-melanophoresabsent on lower jaw until at least 19 mm; cleithral symphysis melanophore absent in most specimens; urostyle pigment present only between about 8 and 11 mm; no pigment on base of PI; no pigment at tips of caudal rays; no pigment along dorsal-fin base anterior to posterior cluster of melanophores until approximately 14 mm. Remarks.-The series is reasonably complete, but only seven specimens larger than 13 mm are available. Sixteen of the 68 specimens examined have a melanophore at the cleithral symphysis. These are from almost throughout the geographical range of the available material (Hawaii, East China Sea, Bismark Sea, western Tasman Sea, Indian Ocean off Sumatra) and range in size from 3.5 to 10.9 mm. Other than this single melanophore, these 16 specimens are indistinguishable from the other P. filamentosus specimens (Table 15, Figs. 6, 30, 31). The distribution of the specimens with the cleithral melanophore includes Hawaii, and the largest three specimens with it are scaled and have 62-64 lateral line scales. This combination of distribution and scale counts eliminates all possibilities within Aphareus or Pristipomoides except P. filamentosus, or less likely, P. zonatus. It is possible that the larvae with the cleithral melanophore represent P. zonatus (although the scale counts are a bit low), but with only this single pigment character to separate the two types, it is equally possible they represent variation within P. filamentosus that is not related to geography. This will be resolved only when the larvae of P. zonatus are identified. P. filamentosus specimens with cleithral symphysis pigment are likely to be confused with Aphareus rutilans larvae prior to the formation of scales. These two taxa can be distinguished because in P. filamentosus larvae of these sizes the only pigment on the P2 is found distally on rays 1 and 2, and there is marked internal structure in the fin spines.

Whereas A. rutilans larvae have pigment on P2 rays 3 and 4 from 7 mm, and lack marked internal spine structure until well after the scales are formed.

Pristipomoides freemani? Anderson, 1966 1 specimen 11.1 mm Figs. 24, 25, 32, Table 16

Material Examined.-Westem Atlantic, vicinity of Florida Keys, NMFSM CA 8910-5108. Diagnosis.-Western Atlantic distribution. MORPHOLOGY-finspines moderately long (Fig. 24); P2sp > Dsp2; moderate internal structure present in fin spines; number of spines on the outer border of the preopercle high (Fig. 25); 52 lateral- line scales; moderate to high number of gill rakers (Fig. 7). PIGMENT-melanophores present on forebrain; melanophores on tip of lower jaw; cleithral melanophore absent; urostyle pigment present; no pigment on base of PI; melanophore series on spines of pelvic fins; caudal rays broken; no pigment anteriorly along base of dorsal-fin soft rays.

Figure 36 (lower). Larvae of Pristipomoides multidens and/or typus. Development of spines on the preopercular outer border (does not include spine at angle). Not plotted are values for six 16.7-46.7 mm specimens of 23-50 spines, respectively. 106 BULLETIN OF MARINE SCIENCE, VOL. 55, NO.1, 1994

Table 17. Selected morphometric and meristic characters of Pristipomoides multidens and/or typus larvae. Measurements in mm. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus.

Pre- Orbit Body Body Pee- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal toral LL 10 Sb length length length length eler length (P) (A) fin fin fin scales spines spines Flexion 4.10 2.35 1.60 1.50 0.58 0.45 1.15 1.00 VII 0 0 4.35 2.51 1.85 1.76 0.66 0.61 1.38 1.08 IX,12 II,7 0 0 4.77 2.70 1.94 1.87 d d 1.55 1.10 IX,12 II,9 0 0 4.83 2.64 1.81 1.74 0.63 0.56 1.34 1.05 Vm,lO 1,7 0 0 4.84 2.64 1.68 1.66 0.59 0.56 1.63 1.16 X,ll H,9 0 0 Postflexion 5.18 3.00 2.11 2.17 0.73 0.69 1.84 1.43 X,11 m,8 -13 0 I 0 5.68 3.23 2.13 2.00 0.78 0.59 1.84 1.38 X,II m,8 ~7 0 I 0 5.68 3.23 2.26 2.13 0.77 0.58 1.94 1.42 X,II Ill,8 ~6 0 I 0 5.92 3.65 2.34 2.41 0.78 0.84 2.00 1.40 X,II Ill,8 -8 0 d d 6.08 3.55 2.45 2.40 0.82 0.84 2.02 1.49 X,II Ill,8 -10 0 1 0 6.45 3.71 2.77 2.68 0.94 0.77 2.19 1.68 X,II III,8 -10 i 1 0 6.67 4.00 2.69 2.69 1.02 0.86 2.31 1.67 X,II III,8 12 0 1 0 6.90 3.93 2.58 2.45 1.00 0.70 2.32 1.74 X,II m,8 14 ] I 7.52 4.72 2.96 2.96 0.96 0.88 2.56 2.00 X,II III, 8 13 2 0 7.55 4.45 2.90 2.71 0.97 0.71 2.58 1.94 X,II III,8 12 2 I 7.81 4.48 2.96 3.08 1.03 0.75 2.77 2.19 X,II III,8 13 3 I 7.83 4.85 3.15 3.00 1.10 0.80 2.66 2.13 X,l] III,8 13 3 1 7.94 4.74 3.07 3.19 1.10 0.81 2.71 2.06 X,ll III,8 12.5 2 I 8.07 4.87 3.20 3.42 1.16 0.90 2.77 2.00 X,II III,8 14 2 1 8.26 4.81 3.10 3.20 1.05 0.81 2.84 2.16 X,l1 Ill,8 13.5 3 I 8.75 5.75 3.75 3.50 1.30 1.20 3.15 2.44 X,I] Ill,8 11 48 3 1 8.94 6.00 3.45 3.25 1.20 1.06 3.25 2.75 X,II III, 8 12 -48 2 I 9.25 5.92 3.70 3.60 1.28 1.20 3.45 2.66 X,II Ill,8 14 48 3 I 9.48 5.80 3.71 3.59 1.28 0.96 3.46 2.69 X,II Ill,8 13 -48 4 I 9.48 5.89 3.71 4.10 1.28 0.90 3.46 2.63 X,ll Ill,8 14 48 4 I 10.38 6.41 3.84 d 1.41 d 3.59 2.69 X,II Ill,8 16 49 2 I 10.57 6.66 4.10 4.04 1.40 1.02 3.59 2.75 X,II Ill,8 16 49 2 I 11.14 6.79 4.36 3.84 1.47 0.96 3.46 2.82 X,II III, 8 IS -49 4 I 11.14 6.66 4.23 4.36 1.47 1.09 3.71 2.88 X, 11 III, 8 -IS -48 5 I 11.79 7.05 4.61 4.48 1.54 1.15 4.04 3.46 X,II III, 8 15.5 50 4 I 12.25 7.75 4.75 4.95 1.66 1.48 4.50 3.90 X,II Ill, 8 IS 48 4 I 12.63 8.63 5.33 4.95 1.59 1.50 4.60 3.80 X,II III, 8 IS 49 4 2 13.58 8.25 5.06 5.25 1.67 1.50 4.87 4.36 X,l1 Ill,8 16 -48 5 2 15.38 9.50 6.25 5.54 2.16 1.72 5.42 4.50 X,II Ill,8 IS 49 6 2 16.70 10.50 6.28 6.53 2.05 1.79 5.25 4.36 X,I] Ill,8 16 53 7 5 18.19 11.27 6.80 6.70 2.30 1.67 6.15 5.51 X,II Ill,8 16 -50 5 4 18.96 11.22 6.97 7.00 2.20 1.74 6.34 5.51 X,II Ill,8 15 49 6.5 4 30.55 17.42 10.45 10.06 3.00 3.00 9.74 8.26 X,II III, 8 16 51 7 6 31.30 18.70 10.50 9.18 3.77 3.12 10.17 8.69 X,II III, 8 16 51 9 6 46.70 26.80 16.20 15.50 5.60 4.20 15.25 13.80 X,II III, 8 16 51 10 12

Remarks.- The identification of the sole specimen is questionable. It is considered a Pristipomoides because it possesses all the characters of the genus (and Aphare- us is absent in the Atlantic). However, it differs from all other Pristipomoides and Aphareus larvae in possessing a series of melanophores in the chevron groove of the spine of the pelvic fin. These melanophores extend from near the base of the spine (Fig. 32). The left spine has three melanophores, and the right has seven. Aside from the P2SP pigment, the specimen differs from P. aquilonaris only in having 16 lower limb gill rakers (P. aquilonaris of that size have 13-14, Fig. 7). This implies an identification of P. freemani, but without a series of larvae, the identification is open to doubt. Further doubt is introduced because, according to LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 107

Figure 37. Larvae of Pristipomoides sieboldii. Scale bars = I mm. A. 3.8 mm (AMS 1.26292-001) from the Philippine Sea. B. 4.2 mm (AMS 1.26142-001) from the western Tasman Sea.

Allen (1985), P. freemani adults do not occur north of Panama. However, even assuming this is correct, it is conceivable that older larvae could be advected as far north as the Florida Keys. Clearly, more larvae are required for study. Because only one specimen is available, most aspects of development are uncertain.

Pristipomoides multidens (Day, 1870) and/or typus Bleeker, 1852 40 specimens, 4.1-46.7 mm Figures 33-36, Table 17

Material Examined.-Gulf of Thailand, USNM Naga S9-11 60-896: Makassar Strait, ZMUC DANA 3796-11, DANA 3796-IV: Pacific Ocean north of Irian Jaya, ZMUC DANA 3768-XVI: Bismarck Sea, AMS 1.19714-015, 1.19726-016: Gulf of Papua, AMS I. 24552-005: Tasman Sea, AMS 1.17875-014, 1.26091-003,1.26141-001,1.26151-003: Indian Ocean over northwest shelf of Australia, AMS 1.26410- 001,1.26411-001,1.26411-002,1.26412-001,1.26413-001; CSIRO ASOI/83 LF Stn.3(1), AS01l83 LF Stn.3(2), AS01l83 LF Stn.14(1), ASOI/83 LF Stn.14(3), AS02/83 LF Stn.7(1), AS02/83 LF Stn.12(3), AS06/82 LF Stn.4(1), AS06/82 LF Stn.7(1), AS06/82 LF Stn.7(3), AS06/82 LF Stn.9(1): 108 BULLETIN OF MARINE SCIENCE. VOL. 55, NO. I, 1994

Figure 38. Larvae of Pristipomoides sieboldii. Scale bars = 1 mm. A. 4.5 mm (CSIRO AS05/82 LF Stn. 14(1)) from the eastern Indian Ocean over the northwest continental shelf of Australia. B. 7.2 mm (MUFS 9504) from East China Sea. C. 9.1 mm (AMS 1.32084-003) from the East China Sea. Only the scales along the lateral line are illustrated. LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 109

Figure 39. Larvae of Pristipomoides sieboldii. Scale bars = I mm. Note the melanophores at the tips of the caudal fin lobes. Both specimens are fully scaled, but only a portion of the scales are illustrated, and myomeres are omitted. Weak internal structure in the fin spines is not illustrated. A. 15.0 mm CAMS1.25362-009) from the Pacific Ocean off the Hawaiian island of Oahu. B. 37.3 mm CAMS1.24163-010) from the western Tasman Sea. Note that scales extend over the proximal portion of the caudal-fin rays.

Indian Ocean west of Sumatra, AMS 1.33623-001; ZMUC DANA 3861-I-V, DANA 3868-1, DANA 3869- II, DANA 3876-11,DANA 3881-1, DANA 3885-1, DANA 3885-1Y. Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-fin spines short (Fig. 35);

Dsp < P2sp at all sizes; internal structure present in fin spines after approximately 7.5-9 mm, becoming strong by 17-30 mm; number of spines on the outer border of the preoperc1e low (Fig. 36); subopercular spines form early (Table 4); scales form early (from 6.5 mm); lateral-line scales 48-53 (after ca. 8 mm); few gill rakers (Fig. 6). PIGMENT-forebrain with melanophores after approximately 7 mm; melanophores consistently present on lower jaw after approximately 12 mm; c1eithral symphysis melanophores absent in most specimens; urostyle pigment present in postflexion individuals; no pigment on base of PI; melanophores present at tips of caudal fin after approximately 12 mm; pigment present along the base of anterior dorsal-fin rays after approximately 7 mm. Remarks.- The series of flexion and postflexion larvae is reasonably complete, but no preflexion larvae were available. However, the series includes some of the largest pelagic specimens of Pristipomoides available. At this size, only a few small serrations remain of the extensive larval head spination. It is unclear if this description is based on one or two species. However, the gill-raker data (Fig. 6) suggest that two species may be present: one with few .•..•. m..J •..•# PrlOllpomolde •• Ieboldll Ql 0 Preflexlon C 40 '0. • Flexion (/) • Poetflexlon U 30 • .; • .\ . • • • Qj •• • • ·: • • 11. •••• • 20 •• •• -0 •• .s::. .. 0 Cll o 0 C 10 Ql 0 ..J .•..•. 00 m..J 40 •..•# cQl 30 • Q. • ~ • 8e•• • • (/) •• • • • • • 0 •• • • • • • • iii 20 0 • ., 0 0 0 0 00 "t:I 10 c: ('oj '0 , . . .s::... 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Cll c Ql Body Length (mm) ...J

•

• • • • • • • • • • ••• • • • • P,latlpomoldea au,lcllla P,lalipomoldee aleboldll •• •• •• • o 0 ••• • Poalllexion o Prellexlon • Flexion 00 • .Poatllexlon 4 5 6 7 8 9 10 11 12 13 14 15 16 Body Length (mm)

Figure 40 (upper). Larvae of Pristipomoides sieboldii. Top. Length of pelvic-fin spine (as percent of body length) in relation to body length. Not plotted are values for nine 18.7-53.8 mm specimens of23.8-14.1 %, respectively. Bottom. Length of dorsal-fin spine 2 (as percent of body length) in relation to body length. Not plotted are values for nine 18.7-53.8 mm specimens of 20.1-11.9%, respectively. LEIS AND LEE: ETELlNAE LARVAL DEVELOPMENT III

Table 18. Selected morphometric and meristic characters of Pristipomoides sieboldii larvae. Mea- surements in mm. d, damaged; i, incipient; a, anlage; (A), at anus; (P), at PI base.

Pre· Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal toral LL 10 Sb length length length length eler length (P) (A) fin fin fin scales spines spines Preflexion 3.10 d 1.25 0.93 0.37 d 0.87 d II a 0 0 0 0 3.40 1.70 1.25 l.l0 0.38 0.25 0.85 0.85 IV a 0 0 0 0 3.65 1.88 1.44 1.22 0.41 0.41 0.97 0.84 IV a 0 0 0 0 3.75 1.85 1.30 l.l0 0.40 0.30 0.93 0.62 II a 0 0 0 0 3.80 2.02 1.48 1.44 0.48 0.46 0.98 0.78 V a 0 0 0 0 3.87 2.13 1.48 1.25 0.44 0.34 0.91 0.78 IV a 0 0 0 0 Flexion 4.19 2.16 1.50 1.38 0.47 0.44 1.25 1.03 VIII i 0 0 0 4.41 2.26 1.71 1.52 0.56 0.47 1.19 1.00 VI i 0 0 0 4.52 2.23 1.68 1.45 0.56 0.44 1.34 1.03 IX 1,8 0 0 4.66 2.53 1.82 1.71 0.60 0.49 1.37 1.05 VII 11,9 0 0 4.89 2.60 1.86 1.75 0.60 0.51 1.47 1.07 IX,12 11,9 0 0 4.98 2.71 1.83 1.64 0.59 0.43 1.36 d IX,I2 d 0 0 Postflexion 5.10 2.79 2.07 1.84 0.64 0.52 1.48 1.26 X, 11 III,8 0 I 0 5.37 2.90 1.83 1.95 0.75 0.51 1.65 1.31 X,II III,8 i 0 I 0 5.38 2.96 2.10 2.08 0.77 0.59 1.86 1.35 X,11 III,8 6 0 1 0 5.61 3.15 2.45 2.13 0.75 0.64 1.71 1.36 X,ll III,8 4 0 1 0 5.86 3.20 2.19 2.20 0.75 0.68 1.77 1.46 X,II III,8 8 0 I 0 5.90 3.19 2.15 2.20 0.77 0.61 1.69 1.35 X,ll III,8 6 0 1 0 6.18 3.66 2.57 2.22 0.80 0.60 1.94 1.54 X,II III,8 5 0 I 0 6.34 3.27 2.41 2.26 0.82 d 1.82 1.45 X,ll III,8 5 0 1 0 6.51 3.48 2.26 2.45 0.84 0.68 2.19 1.68 X,11 III,8 10 0 1 0 6.66 3.85 2.86 2.48 0.89 0.68 2.10 1.75 X,ll III,8 9 0 1 0 6.84 3.77 2.85 2.47 0.82 0.73 2.21 1.74 X,II III,8 13 I 0 6.90 4.00 2.77 2.80 d d 2.13 1.60 X,11 III,8 -11 0 I 0 6.97 3.87 2.71 2.45 0.84 0.71 2.23 1.74 X,II III,8 -10 0 I 0 7.20 4.08 2.64 2.48 0.84 0.76 2.24 1.60 X,ll III,8 12 3 0 7.78 4.11 2.95 2.50 0.89 0.70 2.22 1.69 X,ll III,8 12 I 0 8.12 4.30 2.92 2.81 1.04 0.79 2.30 1.96 X,II III,8 16 2 0 8.48 4.31 3.64 3.14 0.98 0.87 2.55 2.20 X,II III,8 17 i 2 0 8.90 d 3.20 3.17 1.06 0.80 2.51 d X,II III,8 d d 2 0 9.12 5.15 3.20 3.28 1.04 0.88 2.64 2.32 X, 11 III,8 16 72 1.5 0 9.34 d 3.76 3.21 1.02 0.86 2.79 2.27 X,11 III,8 -15 d 2 0 9.60 5.24 3.31 3.33 l.ll 0.80 2.83 2.36 X,II III,8 16 -70 2 0 9.63 5.89 3.49 3.32 1.16 0.80 2.80 2.44 X,11 III,8 16 d 2.5 0 13.96 7.62 4.93 4.90 1.45 1.29 3.93 3.29 X,ll III,8 17 72 3 2 14.10 8.64 5.08 4.59 1.68 1.32 4.16 3.40 X,ll III,8 17 72 3 I 15.00 8.77 5.12 5.06 1.60 1.39 4.26 3.61 X,II III,8 17 70 3 2 18.70 10.63 5.89 5.89 1.79 1.35 4.74 3.97 X,ll III,8 17 -70 d 4 25.80 15.00 8.58 8.07 2.69 2.56 6.53 5.51 X,II III,8 17 70.5 5 5 26.20 14.35 8.71 8.07 2.56 2.05 6.66 5.89 X,ll III,8 16 -68 5 4 37.30 20.50 11.40 11.30 3.00 3.40 8.80 7.90 X,I2 III,8 17 69 5 7 43.70 26.50 14.70 12.70 4.00 3.50 9.60 8.58 X,II III,8 17 72 II 6 44.00 24.50 14.80 14.00 4.10 3.60 10.70 10.00 X,II III,8 I7 69-70 5 13 52.60 28.80 18.30 16.00 4.40 3.80 12.30 10.80 X,ll III,8 17 71 8 14 52.60 30.00 19.40 16.80 4.60 4.50 12.20 10.30 X,II III,8 16 70 7 10 53.80 31.40 19.50 17.60 5.00 5.00 11.90 11.60 X,II III,8 I7 70 9 10

Figure 41 (lower). Larvae of Pristipomoides sieboldii and Pristipomoides auricUla. Development of spines on the preopercular outer border (does not include spine at angle). Not plotted are values for nine 18.7-53.8 mm specimens of P. sieboldii of 18-41 spines, respectively. 112 BULLETIN OF MARINE SCIENCE. VOL. 55. NO. I. 1994

Figure 42. Larvae of Pristipomoides sp. P from (A, C) the western Tasman Sea, and (B, D) the East China Sea. Note melanophore at lower edge of PI base, and multiple melanophores at cleithral symphysis and tip of lower jaw. Scale bars = 1 mm. A. 3.3 mm, head damaged in this specimen (AMS 1.26123- 002). B. 4.4 mrn (MUFS 9509). C. 4.9 mm (AMS 1.3138]-002). Note that there are 12 dorsal and 9 anal fin rays, the first of which are beginning to transform into spines. D. 6.9 mrn (AMS 1.32028-001). LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 113

Table 19. Selected morphometric and meristic characters of Pristipomoides sp. P larvae. Measure- ments in mm. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus. Scales are absent as are spines on the suboperc1e.

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal toral 10 length length length length clef length (P) (A) fin fin fin spines Preflexion 3.25 1.57 1.25 1.03 0.34 0.30 0.80 0.68 III a 0 0 4.24 1.98 1.45 1.44 0.55 0.37 1.11 0.91 VII 0 I Flexion 3.80 1.88 1.57 1.42 0.52 0.40 1.15 0.92 IV a 0 0 4.22 2.24 1.69 1.64 0.51 0.48 1.17 0.91 VI a i I 4.37 2.17 1.74 1.50 0.54 0.42 1.19 0.91 VII i 0 1 4.64 2.33 1.80 1.69 0.58 0.48 1.29 1.00 VIII 11,9 0 I 4.65 2.29 1.74 1.54 0.59 0.46 1.24 0.92 VI i I 4.73 2.28 1.88 1.65 0.57 0.47 1.29 0.94 IX,12 i i 1 4.94 2.32 1.91 1.72 0.60 0.41 1.51 1.13 IX,12 11,9 i I 5.34 2.63 2.04 1.79 0.63 0.56 1.35 1.00 IX,12 11,9 0 1 5.49 3.02 2.08 1.98 0.65 d 1.57 1.25 IX,12 11,9 1 5.61 2.65 2.06 1.66 0.59 0.49 1.53 1.11 X,11 11,9 1 5.80 3.00 2.19 2.05 0.67 0.61 1.54 1.23 X,ll III, 8 1 Postflexion 5.12 2.88 2.04 1.96 0.68 0.60 1.56 1.24 X,11 III,8 5.16 2.95 2.02 1.95 0.66 0.54 1.58 1.17 IX,12 11,9 i 5.39 2.83 2.12 1.92 0.64 0.56 1.59 1.24 X,ll I1I,8 3 6.01 3.39 2.45 2.16 0.76 0.70 1.81 1.38 X,11 III,8 9 6.58 3.42 2.29 2.13 0.71 0.65 1.94 1.42 X,11 I1I,8 8 6.88 3.38 2.24 2.02 0.75 0.65 1.86 1.31 X,11 I1I,7 6 rakers (no more than 11 on the lower limb), and a second with 12 or 13 rakers. Unfortunately, too few larvae are available to resolve this. There is some variability in pigment and its development, and this could be due to the putative series containing more than one species. Pigment is present at the tip of the lower jaw after approximately 12 mm, except that one 8.9 mm specimen has a pair of melanophores on the lower jaw. Four of 17 specimens between 4.4 and 7.9 mm have a melanophore at the cleithral symphysis. Pigment along the base of the soft dorsal fin anteriorly is present in most specimens after approximately 7 mm, but three specimens between 7.4 and 9.3 mm lack this pigment. This anterior cluster of melanophores gradually forms a saddle, as do two clusters that form after approximately 13 mm at the middle of the base of the spiny dorsal fin and on the peduncle (Fig. 34). A fifth saddle forms at the level of the pectoral base. These saddles spread ventrally to form diffuse bars.

Pristipomoides sieboldii (Bleeker, 1857) 45 specimens, 3.1-53.8 mm Figures 37-41, Table 18

Material Examined.-Hawaii, AMS 1.23566-018, 1.24998-009, 1.25347-002, 1.25348-002, 1.25352- 003,1.25362-009.1.25362-020; USNM 322278, 325134: East China Sea, AMS 1.31531-001, 1.31979- 001,1.32037-001,1.32040-001,1.32084-003,1.32087-002; NMST-PL 18, NMST-PL 19; ZMUC DANA 3727-ill: East China Sea, MUFS 9504: Sulu Sea, LACM 36135-4: Philippine Sea, AMS 1.26292-001, 1.31786-001,1.31803-001; NMST-PL 27: Coral Sea, AMS 1.24945-001,1.24949-005: Tasman Sea, AMS 1.24163-009, 1.24163-010, 1.26112-005, 1.26113-007, 1.26142-001: Indian Ocean over northwest shelf of Australia, CSIRO, AS05/82 LF Stn.14(1), AS06/82 Stn.3(2): western Indian Ocean east of Seychelles AMS 1.33151-002: western Indian Ocean north of Madagascar, AMS 1.33053-003, 1.33084- 002, 1.33236-002: western Indian Ocean off South Africa, RUSI 40878. 114 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994 -eft ....J m 40 CD -c: '5. 30 A en AA AA A 0 A 'S; A Q) A A A 20 A a. A A -0 ~ 10 C) -c: CD ....J

-~ ....J a:l 40 -c:CD '5. 30 en & A A AA as 4A A •...rtJ A 0 20 A C 't:J c: C\I 10 0 A - Pristipomoides species P ~ 01 , -c: I CD ....J 3 4 5 6 7 8 9 10 Body Length (mm)

Pristipomoides species P rn 8 CD c: '0. Cf) •... 6 t'I:I '3 0 A ... A A (I) 4 c. A 0 A A.M. (I) ... A a. 2 &A 4A A A A

, , 3 4 5 6 7 8 9

Body Length (mm) LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 115

Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-fin spines moderate in length (Fig. 40); Dsp2 > P2sp until approximately 6 mm; internal structure present in fin spines after approximately 9.5 mm, remaining weak until> 14 mm; number of spines on outer border of preopercle low (Fig. 41); subopercular spines form late (Table 4); scales forming early (after 6.8 mm); lateral-line scales 69-72 (after ca. 9 mm); many gill rakers (Fig. 6). PIGMENT-forebrain pigment forms between 10 and 14 mm; a single melanophore at tip of lower jaw; a melanophore at cleithral symphysis disappears between 7 and 9 mm; internal pigment present at urostyle after approximately 5.4 mm, centered over urostyle in most specimens; no pigment on PI base; melanophores form at tips of caudal rays between 10 and 14 mm; only posterior pigment along dorsal-fin base until between 10 and 14 mm. Remarks.-The series has a gap between 9.6 and 14 mm, and although it extends to 53.8 mm, only nine specimens are larger than 15 mm (Table 18). P. sieboldii is the only lutjanid to occur in both the Indo-Pacific (widely distributed) and the Atlantic (only Vema Seamount, Anderson, 1987). However, it is considered here to be an Indo-Pacific species due to its very limited, southern Atlantic distribution. The larval type identified below as Pristipomoides species S is very similar to P. sieboldii (see remarks under that species). In P. sieboldii the pigment along the base of the dorsal fin spreads anteriorly from the posterior base, in contrast to Pristipomoides sp. S in which a detached, anterior cluster of melanophores forms near the base of the first two soft rays. Smaller larvae are most readily distinguished by the fact that most Pristipomoides sp. S have multiple lower-jaw melanophores while most (87%) small P. sieboldii have only a single melanophore at the tip of the lower jaw (Table 3). Larvae of P. sieboldii are remarkably similar to Aphareus rutilans including the lateral-line scale count and most aspects of pigment in specimens larger than 12 mm. Small P. sieboldii have a cleithral melanophore whereas small A. rutilans do not. In addition, in small P. sieboldii, the center of the melanophore at the dorsal-fin base is posterior to the center of the corresponding anal-fin melanophore, whereas in A. rutilans, the dorsal melanophore is directly above the anal in most specimens. In larger larvae, the development of non-distal pigment on the P2 membrane and rays 3-5, the shorter ascending process of the premaxilla (Fig. 8), and higher number of gill rakers of A. rutilans (Fig. 6) will serve to distinguish the two species (Tables 2, 3).

Pristipomoides species P 19 specimens, 3.3-6.9 mm Figures 42-44, Table 19

Material Examined.-East China Sea, AMS 1.31530-001,1.31804-001,1.32028-001; MUFS 9508, 9509, 9510: western Tasman Sea, AMS 1.26121-005, 1.26123-002, 1.26169-001, 1.28743-025, 1.31381-002,1.32703-001: western Indian Ocean, AMS 1.33068-002.

Figure 43 (upper). Larvae of Pristipomoides sp. P. Top. Length of pelvic-fin spine (as percent of body length) in relation to body length. Bottom. Length of dorsal-fin spine 2 (as percent of body length) in relation to body length. Figure 44 (lower). Larvae of Pristipomoides sp. P. Development of spines on the preopercular outer border (does not include spine at angle). 116 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

Figure 45. Larvae of Prislipomoides sp. S, both from the East China Sea. Inset shows ventral view of lower jaw. Scale bars = I mm. A. 3.4 mm (ZMUC DANA 3723-V-50). B. 5.1 mm (AMS 1.31801- 001).

Diagnosis (? indicates questionable because of limited range of sizes avai/- able).-Indo-Pacific distribution. MORPHOLOGY-Dsp2 moderately long (Fig.

43), Dsp2 > P2sp; no internal structure in fin spines?; spines on outer border of the preopercle few, especially on upper limb (Fig. 44); number of lateral line scales and gill rakers unknown, PIGMENT-no pigment on forebrain?; melanophore ventrally on PI base (often on one side only); one to several melanophores on tip of lower jaw; one to four melanophores at cleithral symphysis; melanophore at posterior end of anal fin slightly anterior to its equivalent dorsal-fin melanophore; urosty1e pigment present after approximately 5 mm, centered over urostyle; no pigment on tips of caudal rays? Remarks.- The largest specimen has additional melanophores along the base of the anal fin, and a gular melanophore, so it is possible larger specimens will be still more heavily pigmented. As the largest specimen available is only 6.9 mm LEIS AND LEE: ETELlNAE LARVAL DEVELOPMENT 117

Figure 46. Larvae of Pristipomoides sp. S both from the East China Sea. Note melanophores on bases of anterior dorsal fin soft rays. Inset shows ventral view of lower jaw. Scale bars = I mm. A. 6.3 mm CAMS1.31816-(01). B. 10.3 mm CAMS1.31810-001). Specimen is fully scaled, but only the scales along the lateral line are illustrated. and no scales are present and the gill raker counts are not definitive (and very difficult to make accurately), it is not possible to provide an identification. Spe- cies P is most similar to P. sieboldii and Pristipomoides sp. S, but has additional pigment in several locations, most notably the PI base, cleithral symphysis (13 of 19 specimens have >1 melanophore) and lower jaw (13 of 19 specimens have >I melanophore). Pristipomoides sp. P is possibly a melanistic variant of one of these other two species: additional pigment at cleithral symphysis and tip of lower jaw is found in Pristipomoides sp. S and a very small proportion P. sieboldii individuals, but pigment at the PI base has not been seen in larvae of either species. Only three specimens of Pristipomoides sp. P have just one melanophore at both cleithral symphysis and lower jaw. If species P is a Pristi- pomoides and is distinct from these two species, the distribution and already identified taxa leave only P. argyrogrammicus, and P. zonatus as possibilities tP- -..J ....,r:n 0) .!: a. 40 (f) () • ~ 30 • •• 0) • i • a.# :-. •• • • a. •• • • • • 0 20 8 - 8 8 s: •... 0 0 Pristipomoides S Cl c:: 0 0) 10 0 Preflexion ..J 8 Flexion -tP- Postflex.ion ..J • ....,r:n 40 0) c:: ena. 30 • • ~ -a•• •• to 0 • , •• • • ...rn 88 • 0 20 0 " 'C c:: 0 C\I 10 -0 , s:•... I , Cl c:: 3 4 5 6 7 8 9 10 11 0) ..J Body Length (mm)

10 • • !IJ 9 0) • .!: 8 • • a. • (f) 7 •• • ... • • • as 6 • • • • ~ • • • ()... 5 • • 0) •• a. 4 8 • •- 0 0) • • Pristipomoides S ... 3 • a. 8 2 000 8 ~8 0 Preflexion 8 1 Flexion , Postflexion . • I I 3 4 5 6 7 8 9 10 11 Body Length (mm)

Figure 47 (upper). Larvae of Pristipomoides sp. S. Top. Length of pelvic-fin spine (as percent of body length) in relation to body length. Bottom. Length of dorsal-fin spine 2 (as percent of body length) in relation to body length. Figure 48 (lower). Larvae of Pristipomoides sp. S. Development of spines on the preopercular outer border (does not include spine at angle). LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 119

(the latter species has not yet been recorded from the western Tasman sea, but could be expected to occur there). There is also the possibility it is Aphareus furca, but until specimens with scales are found, this cannot be resolved. Ad- ditional material is clearly required.

Pristipomoides species S (Aphareus furca or Pristipomoides auricilla) 43 specimens, 3.42 mm-lO.3 mm Figures 45-48, Table 20

Material Examined.-East China Sea, AMS 1.31530-003, 1.31801-001, 1.31810-001, 1.31816-001, 1.31823-001, 1.32084-004, 1.32087-003, 1.32089-002, 1.32090-002; NMST-PL 17-002, NMST-PL 20, NMST-PL 21, NMST-PL 22; ZMUC DANA 3723-V-50; MUFS 9503, 9502: Philippine Sea, near Okinawa, AMS 1.31803-002, 1.31815-001: Makassar Strait, ZMUC DANA 3796-II-20: western Tas- man Sea, AMS 1.26121-005: Indian Ocean, west of Sumatra, ZMUC DANA 3875-II-300. Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-finspines moderately long (Fig. 47), Dsp2 > P2sp until approximately 6.3 mm; internal structure present in spines after approximately 8 mm, but remains very weak; number of spines on outer border of preopercle low (Fig. 48); subopercular spines form early (Table 4); scales form early (from 7.6 mm); 69-73 LL scales (from 7.6 mm); gill rakers moderate in number (Fig. 6); ascending process of premaxilla moderately long (Fig. 8). PIGMENT-pigment on forebrain only in 8.6 and 10.3 mm specimens; usually (79% of specimens) 2 or more melanophores ventrally on lower jaw; melanophore (often more than one) present at symphysis of cleithrum until at least 6.7 mm, and may be present until 8.5 mm; internal pigment on urostyle (usually both dorsally and ventrally) after approximately 5.5 mm; no pigment on the PI base; no pigment on tips of caudal rays; melanophore(s) present anteriorly on base of soft dorsal fin after approximately 6 mm (largest without is 6.9 mm), these may be present on one side only; pigment at posterior base of dorsal fin usually (95% of specimens) posterior to equivalent pigment on anal fin base. Remarks.- This species is very similar to P. sieboldii, differing only in having much earlier development of pigment along the base of the soft dorsal fin anteriorly, and in having multiple melanophores at the symphysis of the lower jaw and cleithral symphysis in many specimens. Two other possible differences are earlier development of scales in species S (6.4 mm) than in sieboldii (7.0 mm), and earlier disappearance of the melanophore on the ventral midline of the caudal peduncle in species S (largest specimen with it 6.6 mm) than in sieboldii (largest specimen with it 8.1 mm), although the smallest specimens lacking this melanophore are about 5 mm in both types of larvae. Among smaller specimens that lack the anterior pigment on the base of the soft dorsal fin, only two had only one melanophore on the lower jaw, but both had more than one melanophore at the cleithral symphysis. However, it is possible that some specimens may have only one melanophore in each location. In this case, confusion will result between P. sieboldii and Pristipomoides sp. S. The vast majority of the specimens of this type of larvae came from the outer edge of the East China Sea. This raises the possibility of geographic variation in pigment. However, as many "normal" P. sieboldii specimens were also taken in this area, it seems unlikely that Pristipomoides sp. S is merely a geographical variant of P. sieboldii. Identification is uncertain, but Pristipomoides sp. S can only be P. auricilla or A. furca if it is not a variant of P. sieboldii. As outlined in the identification section, the most likely identification is A. furca. 120 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

Table 20. Selected morphometric and meristic characters of Pristipomoides species S larvae. Mea- surements in rom. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus.

Pre- Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth Dorsal Anal toral LL 10 Sb length length length length eter length (P) (A) fin fin fin scales spines spines Preflexion 3.42 1.86 1.36 1.34 0.51 0.37 0.98 0.85 IV a 0 0 0 0 3.76 2.00 1.44 1.36 0.52 0.36 1.00 0.84 V a 0 0 0 0 3.93 1.88 1.27 1.31 0.43 0.42 1.07 0.81 IV a 0 0 1 0 Flexion 3.83 2.03 1.52 1.43 0.53 d 1.17 0.95 VI a 0 0 0 0 3.85 2.01 1.48 1.43 0.51 0.39 1.11 0.81 V i 0 0 1 0 4.63 2.29 1.56 1.47 0.56 0.42 1.26 0.96 VB i 0 1 0 4.88 2.62 1.78 1.82 0.63 0.55 1.47 1.21 IX,12 11,9 0 1 0 4.92 2.48 1.78 1.68 0.63 0.46 1.36 1.03 VIII i i 0 1 0 5.43 2.92 1.97 2.05 0.73 0.58 1.60 1.31 X,11 I1I,8 3 0 1 0 5.48 2.81 2.00 2.15 0.72 0.61 1.63 1.21 X,II III,8 3 0 1 0 Postflexion 4.92 2.56 1.81 1.79 0.62 0.52 1.44 1.06 IX,12 d 0 1 0 5.10 2.99 2.16 1.99 0.68 0.68 1.64 1.42 X,l1 III,8 i 0 1 0 5.43 3.23 d d d d 1.64 1.24 X,l1 I1I,8 d 0 1 0 5.72 3.13 2.09 2.09 0.72 0.61 1.64 1.32 X,II III,8 4 0 1 0 5.96 3.19 2.23 2.23 0.74 0.62 1.78 1.46 X,II I1I,8 7 0 I 0 6.00 3.36 2.24 2.12 0.84 0.56 1.88 1.44 X,II III,8 9 0 I 0 6.08 3.39 2.36 2.12 0.78 0.54 1.61 1.38 X,II III,8 4 0 1 0 6.30 3.43 2.32 2.47 0.82 0.68 1.91 1.51 X,l1 III,8 7 0 I 0 6.31 3.56 2.33 2.18 0.81 0.63 1.89 1.48 X,II III,8 4 0 1 0 6.34 3.48 2.28 2.39 0.82 0.68 1.85 1.47 X,II III,8 7 0 1 0 6.45 3.70 2.64 2.36 0.86 0.65 2.08 1.60 X, 11 III,8 12 0 1 0 6.50 3.72 2.76 2.35 0.79 0.72 1.95 1.49 X,ll III,8 9 0 1 0 6.64 3.90 2.69 2.32 0.78 0.67 1.99 1.53 X,II III,8 9 0 1 0 6.71 3.71 2.61 2.46 0.84 0.71 2.03 1.65 X,11 III,8 10 0 2 0 6.87 3.89 2.52 2.60 0.84 0.67 2.08 1.73 X,II III,8 11 0 2 0 7.04 4.48 2.64 2.88 0.92 0.74 2.48 1.83 X,l1 III,8 13 0 2 0 7.39 4.39 3.27 2.86 0.88 0.75 2.17 1.87 X,II III,8 11 0 2 0 7.41 3.97 3.12 2.48 d 0.79 2.52 2.15 X,II III,8 14 0 2 0 7.46 4.03 2.77 2.50 0.87 0.63 2.09 1.71 X,11 III,8 13 0 1 0 7.46 4.09 3.14 2.79 1.01 0.77 2.36 1.86 X,II III,8 13 ~65 3 0 7.62 4.59 3.23 3.00 1.01 0.84 2.43 1.96 X,II III,8 14 0 1 0 7.75 4.33 2.85 2.63 0.90 0.7 2.39 1.86 X,11 III,8 16 -69 1.5 0 8.45 4.89 3.27 2.86 0.98 0.72 2.43 1.96 X,ll III,8 15 -72 2 0 8.47 4.90 3.24 2.82 1.12 0.91 2.82 2.24 X,l1 III,8 16 70 1.5 0 8.60 5.03 3.15 3.15 1.00 O.SS 2.S5 2.20 X,II III,S 16.5 -69.5 2 0 8.62 4.96 3.21 3.21 1.11 0.95 2.58 2.20 X,II III,S 16 -69 2 0 8.92 5.19 3.42 3.01 1.03 0.80 2.69 2.13 X,11 III,8 16 -70 2 1 9.00 5.10 d 3.10 1.00 0.80 2.S0 2.15 X,Il III,S 16 71 2 0 9.07 5.05 3.42 3.17 1.06 0.S4 2.70 2.10 X,11 III,S 16 -71.5 2 0 9.73 5.27 3.72 3.24 1.07 0.91 2.S5 2.37 X,II III,S 16.5 72 2 0 9.73 5.30 3.90 3.34 1.06 1.09 2.94 2.39 X,II III,S 16 69 2 1 10.17 d 3.91 3.45 1.16 0.96 2.92 2.29 X,l1 III,S 16 71.5 3 0 10.30 5.30 3.64 3.60 1.24 0.92 3.12 2.56 X,l1 III,S 16 73 2 1

Unidentified Etelinae larvae two similar species, probably of the same genus Diagnosis.-Indo-Pacific distribution. MORPHOLOGy-fin spines unornamented either internally or externally; dorsal-fin spines short (Dsp2, 20-27% BL); P2sp longest spine, followed by Dsp2 and Dsp3; size at flexion varies with species; LEIS AND LEE: ETELINAE LARVAL DEVELOPMENT 121

Figure 49. Larvae of unidentified Etelinae species 1. Scale bars = 1 mm. A. 3.8 mm from the East China Sea (AMS 1.31824-002). B. 4.9 mm from the western Tasman Sea (AMS 1.26151-002).

PAsp short (3-7% BL); head spines generally small and few in number; scale formation unknown. PIGMENT-no pigment on lower jaw; no dorsal pigment on trunk or tail; only one ventral melanophore, that at posterior of anal-fin base; pigment on forebrain, if present, limited to one ventral melanophore; 0-1 melanophores dorsally on midbrain; no other head pigment; no urostyle pigment; little or no pigment on fins; no pigment at c1eithral symphysis; no pigment centrally on dorsal surface of gas bladder; a band of internal melanophores more or less parallel to c1eithrum from anterior surface of gas bladder to urohyal. 122 BULLETIN OF MARINE SCIENCE, VOL. 55, NO. I, 1994

Table 21. Selected morphometric and meristic characters of larvae of Unidentified Etelinae sp. I. Measurements in men. d, damaged; i, incipient; a, anlage; (P), at PI base; (A), at anus, Scales are absent as are spines on the subopercle.

Pre~ Orbit Body Body Pec- Body Preanal dorsal Head diam- Snout depth depth D'p2 P,'p Dorsal Anal toral PO 10 length length length length eter length (P) (A) length length fin fin fin spines spines Flexion 3.80 2.12 1.52 1.48 0.50 0.40 1.28 1.02 0.86 d VIII 2 PostfIexion 4.16 2.46 1.58 1.58 0.56 0.47 1.49 1.15 1.09 1.10 X,II III,8 3 4.85 2.80 1.70 1.80 0.63 0.45 1.80 1.45 1.33 1.43 X,II III,8 2 3

Description.-MoRPHOLOGY-Body depth moderate, 30-37% BL. Snout short, profile initially concave, becoming smoothly rounded. Mouth large, at most just reaching anterior edge of pupil. Head spines short, PAsp 3-7% BL. 1-2 spines on each preopercular limb. Sb O. 10 1. Op 1 forming between 3.8 and 4.2 mm. Sci 1. Pt O. Notochord flexion begins at 3.8-4.9 mm, depending on species. All fin spines smooth, without obvious internal or external structure. All elements of dorsal and anal fins present during flexion. Dsp2 second longest fin spine.

P2sp longest fin spine. All elements of P2 and A fins present during flexion. Pigment.-Lightly pigmented. Ventrally on tail a single melanophore at posterior end of anal-fin base. Conspicuous internal band of melanophores more-or- less parallel to cleithrum extending from anterior face of gas bladder to PI base in smallest specimens, and on to urohyal in largest specimens. No pigment on dorsal surface of gas bladder; pigment present on posterior face of the gas bladder in one species. Pigment absent on dorsal surface of head, trunk or tail except a single midbrain melanophore in one species. One specimen with a single ventral melanophore on forebrain. Head otherwise without pigment, and no melanophore at c1eithral symphysis. Fins unpigmented except for single, small melanophore distally in chevron of Dsp2 of one specimen. Remarks.-Because only five specimens of 3.8-5.8 mm are available, the description given here is limited and identification remains uncertain. It is likely, for example, that the larvae become more heavily pigmented as they grow. These larvae are most similar to those of Etelis in spination and absence of dorsal pigment on the tail, but differ from them in the very limited pigment on mid- and forebrain, the short Dsp2 relative to the P2sp, and the lack of melanophore series in the chevron groove of the longer fin spines. The differences between the two putative species are slight, but of similar magnitude to the differences among congeners in Pristipomoides and Etelis, however it is possible that study of additional material will reveal that only one species is involved. One possibility is that these larvae represent Pristipomoides argyrogrammicus or P. zonatus. These species have frequently been placed in a separate genus (incorrectly called Tropodinius, see Anderson, 1987).

Unidentified Etelinae species 1 3 specimens, 3.8-4.9 mm Figure 49, Table 21

Material Examined.-East China Sea, AMS 1.31824-002: western Coral Sea, AMS 1.32519-001: western Tasman Sea, AMS 1.26151-002. LETS AND LEE: ETELTNAE LARVAL DEVELOPMENT 123

Figure 50. Larvae of unidentified Etelinae species 2 from the East China Sea. Scale bars = I mm. A. 4.9 mm CAMS1.31530-002). Note that the tenth dorsal-fin element is transforming from a ray to a spine, but that the third anal-fin element is still a ray; dorsal melanophore on the midbrain is on the midline; left maxilla and premaxilla damaged, drawing based on right maxilla and premaxilla. B. 5.7 mm CAMS 1.32042-001). Note that snout, premaxilla and maxilla are damaged, and that midbrain melanophore is actually on the right side.

Diagnosis.-MoRPHOLOGY-body depth greater (than sp. 2), increasing from 34 to 37% BL; small at flexion, just starting to flex at 3.8 mm, fully flexed before 4.2 mm; Dspl remains short. PIGMENT-no pigment on midbrain; melanophores present on gas bladder anteriorly and posteriorly. Remarks.-This species is the most deep-bodied eteline larvae. Unfortunately, the developmental patterns of other eteline larvae provide no confidence that it will remain so throughout its larval development. 124 BULLETIN OF MARINE SCIENCE, VOL. 55, NO.1, 1994

Table 22. Selected morphometric and meristic characters of larvae of unidentified Etelinae sp. 2. Measurements in mm. i, incipient; (P), at PI base; (A), at anus. Scales are absent as are spines on the suboperc1e.

Pre- Orbit Body Body Pee- Body Preanal dorsal Head diam- Snout depth depth Dsp2 P2 sp Dorsal Anal toral PO 10 length length length length eter length (P) (A) length length fin fin fin spines spines Flexion 4.90 2.60 1.65 1.55 0.55 0.50 1.50 1.05 1.12 1.20 IX,12 n,9 3 Postflexion 5.70 3.36 2.16 2.08 0.72 d 1.96 1.48 1.54 1.54 X,ll m,8 9 4

Unidentified eteline species 2 2 specimens, 4.9-5.7 mm Figure 50, Table 22

Material Examined.-East China Sea, AMS 1.31530-002, I. 32042-001. Diagnosis.-MoRPHOLOGY-body depth moderate, increasing from 31 to 35% BL; large at flexion, just starting to flex at 4.9 mm, fully flexed before 5.7 mm; Dspl becomes long. PIGMENT-a single dorsal melanophore posteriorly on midbrain; melanophores present on gas bladder only anteriorly. Remarks.-It is not clear if a very long Dspl (Fig. 50) is characteristic of this taxon, or if it merely represents individual variation. At 12.2% BL, it is the longest Dsp 1 of any eteline larva. Unfortunately, the head of the larger specimen is damaged. More specimens are clearly needed.

ACKNOWLEDGMENTS

This study could not have proceeded without the generous cooperation of many people who made available the coIlections of larval lutjanids 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. Bertlesen, G. Boehlert, T. A. Clarke, B. B. CoIlette, A. Graham, K. E. Hartel, J. Hirota, Y. Iwatsuki, G. D. Johnson, J.-I. Kojima, P. Last, K. C. Lindeman, M. Miya, K. Matsuura, G. E. McGowan, H. G. Moser, B. C. Mundy, J. Nielsen, M. akiyama, T. Ozawa, J. R. Paxton, B. Rachod, W. J. Richards, R. H. Rosenblatt, M. Seki, T. Saruwatari, H. J. Walker, and R. E. Young. L. A. CoIlins, A. C. KendaIl, K. C. Lindeman and W. J. Richards generously abandoned their plans to describe Atlantic Pristipomoides larvae in favor of this paper. D. J. Bray, S. Bullock, S. E. Reader and T. Trnski ably assisted in the lab. T. Goh assisted with data entry and production of the typescript. C. C. Baldwin and W. J. Richards provided valuable criticism of the manuscript. This project was supported by Australian Research Council grant A19031159, and by the Australian Museum. Our very great thanks to all.

LITERATURECITED

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Lutjanus fulviflamma with the changes of body color pattern. Bull. Fac. Agri., Miyazaki Univ. 37(2): 317-321. 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 46~98 in H. G. Moser, W. J. Richards, 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 pp. Kami, H. 1973. The Pristipomoides (Pisces: Lutjanidae) of Guam with notes on their biology. Micronesica 9(1): 97-117. Kyushin, K., K. Amaoka, K. Nakaya and H. Ida. 1977. Fishes of the Indian Ocean. Japan Marine Fishery Resource Research Center, Tokyo. 392 pp. ---, K. Amaoka, K. Nakaya, H. Ida, Y. Tanino and T. Senta. 1982. Fishes of the South China Sea. Japan Marine Fishery Resource Research Center, Tokyo. 333 pp. 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 pp. --- 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 pp. --- and T. Trnski. 1989. The larvae of Indo-Pacific shorefishes. New South Wales Univ. Press, Sydney and Univ. Hawaii Press, Honolulu: 371 pp. Leviton, A. E., R. H. Gibbs, Jr., E. Heal and C. E. Dawson. 1985. Standards in herpetology and ichthyology: part L Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985: 802-832. Masuda, H., K. Amaoka, C. Araga, T. Uyeno and T. Yoshino. 1984. The fishes of the Japanese Archipelago. Tokai Univ. Press, Tokyo, 2 vols. 437 pp. 370 plates. Okamura, 0., K. Amaoka and E Mitani. 1982. Fishes of the Kyushu-Palau Ridge and Tosa Bay. Japan Fisheries Resource Conservation Association, Tokyo. 435 pp. Okiyama, M., ed. 1988. An atlas of the early stage fishes in Japan. Tokai Univ. Press, Tokyo. 1,154 pp. (in Japanese). ---. 1991. Fish metamorphosis: its definition and possible significance. Pages 36-46 in M. Tanaka, ed. Early development in fishes. Koseisha-Koseikaku, Tokyo in Japanese. Potthoff, T., S. Kelly and L. A. Collins. 1988. Osteological development of the red snapper, Lu- tjanus campechanus (Lutjanidae). Bull. Mar. Sci. 43: 1-40. Rosenblatt, R. H., J. E. McCosker and L Rubinoff. 1972. Indo-west Pacific fishes from the Gulf of Chiriqui, Panama. Contrib. Sci., Los Angeles Cty. Mus. 234: 1-18. Senta, T. and S. M. Tan. 1975. On Pristipomoides multidens and typus (Family Lutjanidae). Jap. J. Ichthyol. 22(2): 68-76. Uyeno, T., K. Matsuura and E. Fuju. 1983. Fishes trawled off Surinam and French Guiana. Japan Marine Fishery Resource Research Center, Tokyo. 511 pp.

DATEACCEPTED:October 7, 1993.

ADDRESS:Section of Fishes, Division of Vertebrate Zoology, The Australian Museum, P.O. Box A285, Sydney South, NSW 2000, Australia.

Larval Development in the Lutjanid Subfamily Etelinae (Pisces): the Genera &lt;I&gt;Aphareus, Aprion, Etelis&lt;/I&gt; and &lt;I

Larval Development in the Lutjanid Subfamily Etelinae (Pisces): the Genera <I>Aphareus, Aprion, Etelis</I> and <I