Larval Development in the Lutjanid Subfamily Apsilinae (Pisces): the Genus Paracaesio

BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

LARVAL DEVELOPMENT IN THE LUTJANID SUBFAMILY APSILINAE (PISCES): THE GENUS PARACAESIO

Jeffrey M. Leis, Suzanne Bullock, Dianne J. Bray and Krysia Lee

ABSTRACT Larval development in the lutjanid subfamily Apsilinae is described and illustrated from specimens captured with plankton nets and midwater trawls in the Indian and Pa- cific Oceans. Series were assembled using body shape, fin-spine morphology, head spination, scale formation and counts, gill-raker counts, and pigment patterns. Identifi- cations (mostly to the genus level) were confirmed from adult characters developed in the largest specimens and from the distributions of the adults. Larvae of the Apsilinae 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,10; A III,8), absence of external ornamentation on fin spines, early forming pigment on the midbrain posteriorly, and early formation of the third anal-fin spine and scales. Of the three Indo-Pacific apsiline genera, Lipocheilus, Paracaesio, and Parapristipomoides, we were able to identify only larvae of Paracaesio. However, within this nominal genus are two distinct larval morphologies (morphs), each with several larval types apparently corresponding to species. Due to incomplete understanding of adult taxonomy and distribution in Paracaesio, it was not possible to identify any of the larval types to species. Paracaesio larvae with about 70 lateral-line scales, a relatively slender body and very long, whip-like dorsal-fin

spine 2 (to 96% body length) and equally long pelvic-fin ray 1 (to 370% P2 spine length), scales that formed between 8 and 11 mm, prominent pigment on the urostyle, and no melanophore at the cleithral symphysis are called Paracaesio morph-hi. These were identified as Paracaesio because, among apsilines, only this genus contains species with about 70 lateral-line scales. Five types of Paracaesio morph-hi larvae, distinguished primarily by pigment patterns and secondarily by minor differences in head spination, are described from larvae as large as 24 mm. For one larval type, apparently of the Paracaesio morph-hi group, the largest specimen (6.2 mm) was too small to have scales. Therefore, we could not establish that it was a Paracaesio species. We describe this very lightly pigmented larval type under the name Unidentified Apsilinae. Its South Pacific distribution nearly matches that of Parapristipomoides squamimaxillaris, so the possibility remains it is that species. Larvae considered to be Paracaesio because they have about 50 lateral-line scales, but too many gill-rakers to be Lipocheilus carnolabrum, are called Paracaesio morph-lo. These larvae have a relatively deep body, relatively short, robust

dorsal-fin spine 2 (to 31% body length) and pelvic-fin ray 1 (to 125% P2 spine length), scales that form between 7 and 9 mm, a melanophore at the cleithral symphysis until 4-7 mm, and no pigment on the urostyle. Three types of larvae of this morph are described from larvae as large as 29 mm, and are distinguished by spination on supraorbital ridge, and lachrymal, and minor pigment characters. In one type, the largest larva was too small for us to be sure it had the adult complement of gill-rakers, and there is a possibility it is L. carnolabrum rather than a Paracaesio species. Another type was identified as either Paracaesio gonzalesi or P. kusakarii based on scalation and pigment. The presence of two, distinct morphs of Paracaesio larvae supports a suggestion made by W. D. Ander- son, Jr., based on adult characteristics, that there are two monophyletic lines within this nominal genus. However, with the possible exception of one pigment character, larvae seem to provide no evidence that Paracaesio is monophyletic.

697 698 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

The lutjanids of the subfamily Apsilinae (sensu Johnson, 1980) are medium-sized fishes found at moderate depths (20-460 m) of oceanic island and continental habitats in sub- tropical and tropical waters of both the Atlantic and Indo-Pacific (Allen, 1985; Anderson, 1987; Anderson et al., 1992). Some species are important in commercial and artisanal hand-line fisheries. The four genera —Apsilus, Lipocheilus, Paracaesio, and Parapristipomoides — are not well known either biologically or taxonomically. The two species of Apsilus are confined to warm Atlantic waters. The monotypic genus Lipocheilus occurs on hard bottoms of the deeper portions of the continental shelf and upper slope from the Arabian and Andaman Seas to the northwest continental shelf of Australia, New Britain, the Philip- pines and the Ryukyus (Allen, 1985; Kailola, 1987; Allen and Swainston, 1988; W. D. Anderson, Jr., pers. comm.). The monotypic genus Parapristipomoides is known from rocky bottoms at 100-400 m in the South Pacific from Rapa and Easter Island in the east (Allen, 1985), Tonga (Randall, et al., 1990) and New Caledonia and the Coral Sea in the west (Rivaton et al., 1989, Kulbicki et al., 1994). Paracaesio spp. are found over rocky bottoms and coral at moderate depths (20-250 m) throughout the warm portions of the Indo-Pacific except Hawaii (Allen, 1985; Randall, 1985; Anderson, et al., 1992), but the number of species and their distributions are unclear. Anderson et al. (1992) recently described two new species of Paracaesio and estimated there are “about 8 or 9 species” in the genus, but mentioned in an addendum the possibility that two additional undescribed species are present in the western Indian Ocean. Anderson (1994) further discussed taxonomic problems in the genus Paracaesio. As the name suggests, Paracaesio bears a general resemblance to some of the caesionine lutjanids, and occupies a similar zooplankton-feeding, schooling niche, albeit in generally deeper water. There are only two published accounts of larvae of apsilines. Leis (1987) illustrated and very briefly described a single 6-mm larva identified only as an apsiline lutjanid. This larva has subsequently been identified as a Paracaesio species. Okiyama (1991) illustrated, as an example of a prejuvenile early life history stage, an 11.2-mm individual identified as “aodai”, the Japanese common name for Paracaesio species. Nothing is known about spawning in apsiline lutjanids, but they presumably produce pelagic eggs as do all lutjanids for which spawning mode is known (Leis, 1987). Our efforts to describe larval development in this subfamily were hampered by limited availability of material. We were unsuccessful in obtaining larvae of the Atlantic Apsilus. If we were successful in obtaining any Lipocheilus or Parapristipomoides larvae, then they were only smaller larvae we have been unable to identify as such. However, for Paracaesio, we were apparently too successful, and have assembled what appear to be more types of larvae than there are accepted species of adults. Furthermore, there are two distinct morphs of larvae attributable to Paracaesio spp. as currently understood. These correspond to and corroborate the two possibly monophyletic groupings of species tentatively recognized by Anderson et al. (1992). A further complication is that distributions of most Paracaesio spp. are incompletely known (Anderson et al., 1992; Kramer et al., 1994). Unfortunately, because of the above, we were able to identify our larval types of Paracaesio only to groups of species, not to individual species. Therefore, this paper contains descriptions of the larval types of Paracaesio to docu- ment the developmental diversity we encountered. The larval development of these species provides information that will shed light on the relationships of the subfamily and LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 699

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Flsyarni LnateraScaleso DlorsalAlnaPsectoraGsillrakerlainescalemaxill Lipocheilus L.carnolabrumX8,10I6II,115-154-7+11-14=17-24o9-5N Paracaesio (Paracaesiomorph-lo) P.caerulea X8,10I6II,1290-12+19-20=28-34o9-5N P.gonzalesiX8,(9)-10I7II,116-1191+18-20=29-34s8-4Ye P.kusakani X8,10I6II,1270-12+19-20=26-34s8-5Ye P.stonei X8,l0I6II,19100-11+17-18=27-24o9-5N (Paracaesiomorph-hi) P.ParagrapsimodonX8,10I7II,1090+20-21=29-36o9-7N P.sordida X9,(9)-10I7II,8-146-192-11+19-24=28-36o9-7N P.waltervadiX8,10I7II,1380-11+21-22=29-36s9-7Ye P.xanthura*X),(9)-10-(11)I8II,8-(9156-172-12+20-23=28-36o8-7YesorN Parapristipomoides P.squamimaxillarisX8,10I6II,16150-12+22-24=32-35s4-5Ye *referredtoasapossiblespeciescomplexbyAndersonetal.(1992). within it. This information will be mentioned only briefly here, as it will be more fully analyzed in a separate paper. These descriptions and the information on larval distribution we provide may also prove helpful in alpha-level taxonomic work in this problematic genus. This paper is the fourth in a series describing larval development in the subfamilies of lutjanid fishes. The others are: Etelinae (Leis and Lee, 1994); Paradicichthyinae (Leis and Bray, 1995); and Caesioninae (Reader and Leis, 1996). A fifth paper on the Lutjaninae is in preparation.

MATERIALS AND METHODS

The genera and species of apsiline lutjanids recognized by Johnson (1980), Anderson (1987) and Anderson et al. (1992) are listed in Table 1. Adult apsiline lutjanids occur in difficult-to-sample, relatively deep habitats, and are most diverse in the Indo-Pacific, where ichthyological exploration is incomplete. So it is likely that some undescribed taxa remain (Anderson, et al. 1992; Anderson, 1994). Distributions of apsiline species are poorly known, so it would not be unexpected to find larvae of a species outside known adult distributions. Among adults, species are distinguished by lateral-line scale counts, whether the maxilla is scaled or not, caudal-fin shape, color, and a few other scale and morphometric characters. Only the first two are of potential help in identifying larvae. Definitions, measurements and abbreviations follow Leis and Trnski (1989) with the exception that the eye diameter is of the orbit, not the pigmented area. Abbreviations used are: A - anal fin;

BL - body length; D - dorsal fin; P1 - 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 700 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997 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 P_O or P_I refer 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; Pcl - postcleithrum; Pe - pterotic. Gill-raker counts on the first arch were made only on larvae >8.0 mm. 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 approxi- mate. 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), very long fin elements, and pigment patterns not found in the bottom-associated juveniles and adults. In some cases, the smallest specimens are not the least developed (in terms of fin formation or development of spines on the head). This is presumably due to differential shrinkage among specimens. In such cases, we may refer to the least developed specimen of a series rather than the smallest specimen. Larvae were examined under a dissecting microscope. Precision of measurements varied with the magnification used: this varied from 8 to 50X 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 EXAMINED. —The material utilized in this study was obtained from a wide variety of institutions. Methods of capture, fixation, preservation, storage and time since capture varied widely, and this has had a marked effect on the condition of the larvae and, hence the detail and complete- ness of the descriptions herein. Many specimens are damaged and the elongate elements of the dorsal and pelvic fins 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 of characters based on spine length and fin pigment, which is unfortunate, as these are important in distinguishing species within genera. 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 ZMUC DANA 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 unless in poor condition. Surplus material is not listed in the “Material Examined”. Institutional abbreviations follow Leviton, et al. (1985), with the addition of MUFS which indicates Miyazaki University Department of Fisheries Science, Miyazaki, Japan, and CBM which indicates Natural History Museum and Institute, Chiba, Japan. A large amount of the material examined was collected by the Ocean Research Institute of the University of Tokyo in a series of cruises between Taiwan and Okinawa, so strictly speaking, some is from the northwest Philippine Sea and some from the eastern edge of the East China Sea. For convenience, all of this material is referred to as having come from the East China Sea. LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 701

tahteton(eavraldinajtulenilispafosepyttnereffidehtgnitarapesniesufosretcarahC.2elbaT foeavral sulispA , suliehcopiL dna, sediomopitsirparaP nevigeraseulaverehW.)nwonknuera .niatrecnu,?;degamad,d.slaudividninoixelf-tsopotreferyeht,][ni

oiseacaraP deifitnedinU oiseacaraP Meih-hpro AbpsilinaMorph- ()Figs.2-6)()Fig.7(Figs.9-13 B]odydepth(%BL)2)3-38[27-382]0-37[31.928-46[38-46 L]engthDsp2(%BL)2]4-96[63-960]-80[55d?10-31[14-29 S,hapeDsp2s,lenderstlenderrobus wehip-likew)hip-lik(Fig.1C,D (Fig.1A,B) SsheathonDsp2y?enoon withpigmented“flag” I;nternalstructurefeormslatef;ormslatformsearly ionDspineswoeakt moderatet mgoderatestron ()Fig.1A,B)(Fig.1C,D

LengthP2r0ay11550-371782-2790-12

(%P2sp) S8?7izeatwhichscales begintoform(mm) S1izebywhichscales1?9 areformed(mm) N0umberofLLscalesa?bout7 about5 0 Moelanophoreatnonlyes,unti cmleithralsymphysis 4-7m Psigmentonurostyleyseyoen M,elanophoreatbaseyoesulsuallynyes,unti osfposterioranalmnostspecie 6mm,the fin-rays usuallyno Psigmentdorsallyyoensye onmidbrain

The Literature Cited section does not include citations based solely on the authorities for the genera and species. These are given by Allen (1985), Anderson (1987) and Anderson et al. (1992).

IDENTIFICATION

All identifications are based on the series method. Because larvae were not identifi- able to species and the number of larval types differs from the number of recognized species, and due to taxonomic uncertainties at the adult level, the possibility remains that some of the putative series (herein called types) contain more than one species. For this reason 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: a corroborating character was required. This approach has the virtue of not over-splitting, but it could lead to over-lumping. 702 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

LUTJANIDAE. Larvae were identified as lutjanids through the characters listed by Leis and Rennis (1983) and Leis (1987) including 24 myomeres; very long dorsal-fin spines (particularly the second spine) and pelvic-fin spine, which are smooth to coarsely serrate but without recurved spinelets; Dsp2 and P2sp first fin elements to form; longest P2 ray similar in length to or longer than P2sp; strong head spination, without serrations on preopercular spines (except some lutjanine and caesionine taxa, and in these, serrations weak); head spination includes weak anterior frontal ridge, supraorbital ridge, weak pterotic ridge, and spines on opercle, subopercle, and interopercle, and on inner and outer borders of preopercle; spines also present on posttemporal, supracleithrum and dorsal postcleithrum; no supraoccipital crest or spines; no lachrymal spines or serrations (except in one apsiline taxon); body and head laterally compressed; and series of ventral melanophores on tail at least in smaller preflexion larvae. APSILINAE. Lutjanid larvae with fin counts strongly modal at DX,10 and AIII,8 considered apsilines, as this combination of modes not found among other lutjanids (Anderson, 1987). Indo-Pacific apsiline larvae further characterized by: external ornamentation on fin spines lacking; internal structure first evident in fin spines either during flexion or shortly thereafter (6 mm); early formation of Asp3 (i.e., during flexion or within 1 mm thereafter); early addition of spines to outer, upper border of preopercle; development of parallel ridges on supratemporal-intertemporal; early development of scales; and early development of dorsal pigment on midbrain (with one exception, see Unidentified Apsilinae, below). PARACAESIO. No apsiline larvae were available from Atlantic which eliminates Apsilus. Parapristipomoides has 54-55 LL scales, and Lipocheilus has 49-54 LL scales, so Indo- Pacific apsiline larvae with about 70 LL scales are Paracaesio (49-72 LL scales, Table 1), and are referred to as Paracaesio morph-hi (Table 2). Parapristipomoides is eliminated among series containing larvae possessing scales, because none of these had 55-58 LL scales. Apsiline larvae with about 50 LL scales might be either Lipocheilus or Paracaesio. The available larvae larger than 11 mm have more gill-rakers than do adult Lipocheilus, and the adult complement of gill-rakers in Paracaesio spp is greater than that in Lipocheilus carnolabrum (Table 1). Therefore, Paracaesio larvae with about 50 LL scales are referred to as Paracaesio morph-lo if their series contains larvae > 11 mm (Table 2) or if they do not occur within the known range of L. carnolabrum. PARACAESIO MORPH-HI. These larvae have about 70 LL scales, moderate body depth, and extremely elongate, slender, whip-like Dsp2. Scales begin to form at about 8 mm, but ossified scales not present on most of body until about 11 mm (Table 2). By morph, we mean a group of morphologically very similar types. We distinguished what appear to be five types of larvae within this morph differing primarily in pigment (Table 3). We were unable to identify any of these larval types to species, but can confidently place them amongst the group of species with an “increased number of lateral-line scales” identified by Anderson et al. (1992): this includes P. paragrapsimodon, P. sordida, P. waltervadi and P. xanthura complex (sensu Anderson, et al., 1992). PARACAESIO MORPH-LO. These larvae have about 50 lateral-line scales, relatively deep body, and relatively short spines in dorsal fin. Scales begin to form at about 7 mm, and full set of scales present before 9 mm (Table 2). We distinguished what appear to be three types of larvae of this morph. One of these larval types has a single, weak spine on the supraorbital ridge. Unfortu- nately, the largest specimen (9.2 mm) cannot be identified further. This type is referred to LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 703

as Paracaesio morph-lo D, and could be any of the four species of Paracaesio with about 50 LL scales. However, the largest specimen is apparently too small to have definitive gill-raker counts, and it is therefore possible, but unlikely, that Paracaesio morph-lo D could be Lipocheilus: see Remarks section under description of this type. The second type of Paracaesio morph-lo larva is represented by only three specimens of about 30 mm. These larvae have a serrate lachrymal, scales on the maxilla, and faint broad bars on the body. They are identified as P. kusakarii and/or P. gonzalesi, the only known Paracaesio species with about 50 lateral-line scales and scales on the maxilla (Anderson et al., 1992). Adults of both species have bars on the body. The other larval type of this morph lacks a spine on the central portion of the supraorbital ridge, scales on the maxilla, and pigmented bars (but the largest is only 20 mm). Within this type are some minor differences in pigment: we were unable to determine if these represent intraspecific variation or the presence of more than one species. However in other lutjanid subfamilies such small differences in pigment often distinguish different species. The three pigment variants (A,B,C), are not described separately, rather a description of the combined type is provided with mention of what constitutes the variable pigment. Because we have distinguished fewer morph-lo types than there are species of Paracaesio with about 50 LL scales, it is likely that the series of this type contains more than one species, regardless of whether the pigment differences are species-specific. This larval type is referred to as Paracaesio morph-lo A-C, and could be any one or more of four species of Paracaesio with about 50 LL scales: P. caerulea, P. stonei, or if pigmented bars and maxillary scales do not form until after 20 mm, P. kusakarii and/or P. gonzalesi. UNIDENTIFIED APSILINAE. This constitutes a partial series of larvae (2.1-6.2 mm) none of which have scales. These larvae are from the South Pacific, outside the documented range of Lipocheilus, but within the known ranges of Paracaesio and Parapristipomoides. These larvae are morphologically very similar to Paracaesio morph-hi larvae, but are very lightly pigmented, and lack pigment on the midbrain (Table 2). It seems likely they are a Paracaesio morph-hi species, but without larger larvae, we cannot eliminate the possibility they are Parapristipomoides, so we refer to them as Unidentified Apsilinae.

DESCRIPTION OF DEVELOPMENT

Paracaesio morph-hi Table 2, Figures 1-6

Diagnosis. —MORPHOLOGY. - (values in [ ] refer to postflexion individuals) - Body moderately deep (23-38[27-38]% BL); head spines moderate in number and size; PAsp reaching maximum of 12.5% BL; Dsp2 very slender and very long (24-96[63-96]% BL), much longer than other Dsp (Dsp3 2-24[15-24]% BL); cross-section of Dsp2 chevron-shape proximally, but round distally; Dsp2 enclosed in distal sheath with small, fleshy “flag” at

its tip; P2sp of moderate length (9-31[20-31]% BL), but P2 ray 1 nearly as long as Dsp2 (150-370% P2sp); scales begin to form at 8-9 mm, but not fully formed until about 11 mm; about 70 LL scales. PIGMENT. - No melanophore at cleithral symphysis; external melanophores on urostyle before flexion, and persisting; external pigment at posterior end of dorsal-fin base in some taxa; “flag” at Dsp2 tip pigmented; postflexion larvae

with extensive pigment along Dsp2 chevron groove and sheath and along P2 rays 1 and 2. 704 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 1. Photomicrographs of internal structure in the spines of the dorsal fin in Paracaesio larvae. Scale bar applies to all photos. Photos by B. Ewart. A. Paracaesio morph-hi type A, 6.24 mm (RUSI 48810), Dsp2 and 3, left side. No internal structure present. Note how spines in illustrations A and B (morph-hi) are much more slender than those in illustrations C and D (morph- lo). B. Paracaesio morph-hi type C, 11.4 mm (ZMA 121.533), Dsp2 and 3, right side. Internal structure is of moderate strength: compare with illustration D. C. Paracaesio morph-lo type A, 6.60 mm (NSMT-PL 41), Dsp2, 3 and 4, left side. Internal structure developing. Note how spines in illustrations C and D (morph-lo) are much more robust than those in illustrations A and B (morph- hi). D. Paracaesio morph-lo, type B, 11.7 mm (AMS I.32063-002), right side. Internal structure is deeper and more distinct than that in morph-hi of similar size: compare with illustration B.

Description. —MORPHOLOGY. -Body compressed and of moderate depth (23-38% BL). The youngest larvae somewhat more elongate, and in postflexion larvae BD 27-38% BL. Body depth increases gradually, with maximum relative depth at about

7-9 mm. Body deeper at P2 base than at anus. Gut, coiled in smallest specimens, also deepens, taking on more triangular shape. Relative position of P2 changes probably due to allometric growth. P2 appears to move from initial position at about level of middle of P1 forward to about level of P1 base (= level of Dsp1). These changes accomplished well before notochord flexion begins. Subsequently, P2 gradually moves posteriorly relative to P1 and D fin to level of Dsp2-4. Small gap between anus and A fin origin disappears shortly after flexion. Head large and compressed. Snout length less than eye width. Snout initially pointed, but as head profile becomes steeper, snout becomes rounded. Mouth large, somewhat oblique and reaches at least anterior margin of eye, but does not quite extend to mid eye. Small, conical, teeth first form anteriorly on premaxillae at about the time Dsp2 forming. Dentary teeth form soon after. Small canine tooth anteriorly in premaxilla forms at about 11 mm. No large canines form. Nasal pit roofs over to form two nostrils at 8-9 mm. LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 705

Spination on head relatively weakly developed for a lutjanid. Head spines smooth and no ridges have serrations (serrations on preopercular outer border seen late in larval stage formed by large increase in number of spines along that border). Least developed specimen (2.65 mm) has no spines on head. Sequence of (and size at) formation of first spine on head bones is preopercle (2.7 mm), interopercle (3.0-3.5 mm), supracleithrum (3.0 mm), posttemporal (3.0-4.0 mm), subopercle (15 mm). Spine at angle of preopercle largest on head, reaching 12.5% BL. It gradually becomes engulfed by preopercle border, but is prominent in largest specimen. Preopercle inner border has 3-4 small spines, all present by 6.5-7.0 mm. Outer border of preopercle armed with spines up to pupil width. First of these at about 3 mm, and spines gradually increase in number. Spine growth does not keep pace with fish, and they eventually form serrate border on preopercle. On upper limb of outer border of preopercle, only one spine present until about end of flexion (5.6-6.2 mm) after which spines added at regular intervals. Interopercular spination initially as small spine just dorsal to and approximately parallel to spine at preopercular angle. Up to four additional small spines added to border of interopercle from 12 mm. Only two largest specimens (15 and 24 mm) have spines on subopercle (2 and 8, respectively). Single opercular spine appears at about 3.5 mm.

A small spine on dorsal postcleithrum just dorsal to P1 base at about 5.0-6.0 mm near completion of flexion, persisting in largest specimen. Supracleithrum develops first spine at about 3.5 mm and adds two more small spines by 5.5-7.0 mm. Small posttemporal spine on dorsal ridge at about 3.5-4.0 mm, and a second spine at 7-8 mm in some species. On ventral ridge of posttemporal a small, single spine forms near completion of flexion. Smooth supraorbital ridge at about 3.5-4.0 mm, persisting in largest individual. A low, smooth frontal ridge dorsal and slightly anterior to supraorbital ridge at about 8 mm. A low smooth pterotic ridge at about 6 mm approximately in line with supraorbital ridge. At about 9 mm, a pair of very low parallel ridges dorsal to posterior end of pterotic ridge: these appear to correspond to the edges of the supratemporal-intertemporal bones (Potthoff et al., 1988). A very small spine may form near dorsal edge of each ridge. These ridges form sides of one sensory canal of head and join dorsally sometime after 15 mm. Extend- ing anteriorly from dorsal end of more anterior of these ridges is very weak ridge that also forms portion of sensory canals. Maxilla develops anteriorly an acute spine-like angle, probably an enlargement of maxillary process. At about 12 mm, lachrymal begins to project over maxilla posteriorly and to cover it when mouth closed. Border of lachrymal smooth to irregular, but lacks spines or serrations. Fin spines and their development are among the most distinctive characteristics of

Paracaesio morph-hi larvae. Least developed specimens have Dsp2 and 3 and P2I,2.

Most of Dsp and P2 rays form before D soft rays, elements of A fin and the caudal rays begin to form. However, latter all present by completion of flexion. Flexion begins at about 4.3-4.5 mm and is complete at about 4.7-5.6 mm. Except for Dsp1, D spines form from anterior to posterior. Ultimate spine of D fin forms initially as soft ray, but remains so over only very short range of BL, and transforms into spine before flexion complete or very shortly thereafter. Asp3 forms initially as soft ray early in flexion stage and transforms into spine during flexion or very shortly thereafter. P1 rays last to form, and all present at about 8.0-8.5 mm, at about time scales begin to form. Dsp2, P2 ray 1 and P2sp become very elongate, especially first two: Dsp2, 24-96[63-96]% BL; P2sp, 9-31[20-

31]% BL; P2 ray 1, 150-370% P2sp length. Dsp2 extremely long and whip-like, and much longer than either Dsp3 (3-24[15-24]% BL) or the P2sp at all times. Maximum relative 706 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

spine length reached between 5 and 9 mm. At least P2 ray 1 and often P2 ray 2 longer than

P2sp, the first exceptionally so (it is about same length as Dsp2). Asp not particularly elongate. All fin spines chevron-shaped in cross-section, at least basally, and concave (trailing) surface contains tissue. This frequently pigmented in Dsp2. Dsp2 chevron- shaped basally: however, majority of spine circular in cross-section and of very small diameter. Extending distally from tissue of chevron groove is a spine-surrounding sheath,

and a small, often pigmented “flag” of tissue at spine tip. On leading edge of P2sp and first Dsp are two ridges basally and a single ridge distally. Basal ridges first obvious between 6-8 mm, and extend about 10-20% (P2sp) or about 50% (Dsp1) length of spine. No fin spines ornamented externally, but distinctive internal structure first develops at 5- 6.5 mm in Dsp2 and 3, and by 7-8 mm in spines of other fins (Fig. 1). It eventually spreads to most of fin spines by about 12 mm. In all but one species (Paracaesio morph-hi type A), gill rakers increase from 17 (3+1+13) at 8 mm to 20 (5+1+14) at 9 mm. At 10 mm, there are 21 (5+1+15) gill rakers, and 23 (6+1+16) between 11 and 15 mm. The 23.5 mm specimen has 25 (7+1+17) gill rakers. In Paracaesio morph-hi type A, there are 18 (5+1+12) gill rakers at 8 mm and 19 (5+1+13) at 14 mm. Morph-hi adults have 28-35 gill rakers (Table 1). Scales begin to form relatively early. Incipient scales present from about 8-9 mm, but full complements of scales not present until about 11 mm. This growth interval presumably means that scales slow to form. No specimens have scales on maxilla. PIGMENT. - Generally light. Pigment cap present dorsally on gas bladder and gut. No melanophore on preanal fin fold. Pigment dorsally on midbrain in all but least developed specimens (< 3 mm). These dorsal melanophores initially located dorso-laterally on posterior portion of midbrain. They increase in number and spread to cover midbrain by about 10 mm. Melanophores absent on forebrain until some unknown size after 15 mm, but in largest specimen (23.5 mm), they cover it. At tip of lower jaw a melanophore may be present or absent (Table 3). An internal melanophore may appear at base of opercular spine from about 10 mm. No melanophore at cleithral symphysis. Initially a series of up to nine small melanophores along ventral midline of tail. These decrease in number so that by end of flexion, at most only three melanophores present, usually one at posterior end of A fin (Table 3) and one or two on caudal peduncle. By 6 mm, only melanophore at A fin base present, and it has become at least half internal if present at all. This melanophore absent in one larval type. Pigment appears at urostyle base very early, first as a melanophore on caudal-fin anlage. This external melanophore appears to migrate to ventral surface of notochord before flexion begins, and thereafter is located over urostyle, and may increase to two melanophores during flexion. By 8 mm, as many as seven external melanophores present along with internal melanophores pro- ducing prominent spot covering urostyle base, although one species retains only a single, small melanophore (Table 3). Pigment on dorsal surface of trunk absent and that on tail limited and varies among taxa (Table 3). D fin pigment associated with the chevron-groove tissue and sheath of Dsp2. Initially, only distal “flag” is pigmented, but in postflexion larvae, a series of melanophores appears, and may extend length of spine. After 10 mm, pigment may also be found on distal parts of membrane of spiny and soft portions of D fin. After about 14 mm, pigment on Dsp2 spreads onto membrane between Dsp2 and 3, and a scattering of melanophores present along base of fin in largest specimen. Only largest specimen has pigment on anal

fin - a distal band on membrane (Fig. 6B). Pigment absent on P1 fins. On caudal fin, LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 707 pigment in most larval types initially limited to a small melanophore initially on anlage and later on base of fin (either dorsally or ventrally), but this generally absent after 8 mm (Table 3). In types A and F, usually more than one melanophore. Some individuals of most morph-hi types (and Unidentified Apsilinae) have one or more small melanophores at base of dorsal caudal rays. After about 10 mm, a few melanophores appear distally on edges of outer caudal rays, both dorsally and ventrally, spreading to form faint bands.

Extent and timing of appearance of pigment on P2 are difficult to describe accurately due to damage to fins of nearly all specimens, but pigment initially confined to edges of P2 ray 1, subsequently spreading to rays 2 and 3. Pigment not present basally, but originates at about level of half the length of P2sp, and probably continues for most of length of rays proximally. Okiyama (1991) illustrated pigmented “flag” at tip of P2 ray 1 in 11.2 mm specimen of Paracaesio morph-hi of unknown type.

Remarks. —The extremely long Dsp2 and P2 ray 1 are fragile and damaged in nearly all our specimens of Paracaesio morph hi. In some cases, the element is broken, but still attached. In others it may be cleanly and obviously broken off, but it is often difficult to determine if all of the element is present. This is particularly true of Dsp2, and one can only be absolutely sure the spine is intact if the pigmented “flag” is still attached. The P2 “flag” illustrated by Okiyama (1991) in an 11.2-mm morph-hi larva is absent in our morph- hi specimens probably due to damage (however, one specimen of Unidentified Apsilinae has this pigmented “flag”, see below). It appears that Dsp2 and P2 are of similar length when intact, giving the larvae a spectacular appearance (e.g., Fig. 6B). Among the different types of Paracaesio morph-hi larvae described here, there are almost no morphological differences, the various types being based on minor differences in pigment (Table 3) and some slight differences in head spination. Unfortunately, because taxonomy and distribution of the adults of this morph are unclear, we cannot be certain that these types constitute distinct species. However, in the eteline lutjanid genus Pristipomoides, morphological differences among types of larvae are also very limited (Leis and Lee, 1994), and the kinds of differences in pigmentation used to distinguish types of Paracaesio morph-hi larvae are reasonably reliable indicators of differences among Pristipomoides species. This suggests that the types described here represent different species, but more larval material and a clearer understanding of Paracaesio taxonomy and distribution will be required to be sure. In fact, biochemical methods may offer the best means of establishing identifications in larvae of this morph.

Paracaesio morph-hi type A 10 specimens, 3.1-14.0 mm Figure 2, Tables 3, 4

Material Examined. - East China Sea, AMS I.32089-008, I.31802-001: Sulu Sea, USNM 337337: western Indian Ocean, AMS I.33135-004, I.33161-004, I.33221-003; RUSI 48810, 48811.

Diagnosis. —MORPHOLOGY - morph-hi; second spine on dorsal ridge of Pt absent; third spine of Scl forms very late (8 mm). PIGMENT. - No dorsal pigment on trunk or tail. Melanophore present at tip of lower jaw. No melanophore at base of A fin posteriorly. Usually more than one melanophore at base of caudal rays. 708 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 2. Larva of Paracaesio morph-hi type A, 6.24 mm from the western Indian Ocean off South Africa (RUSI 48810). Dsp2 is broken, the third element of the anal fin is transforming from a soft ray to a spine, and none of the fin spines have any internal structure. Scale bar is 1 mm.

Remarks. —Few specimens of this type are available. Type A is characterized by sparse pigment, most obviously the lack of a melanophore at the base of the anal fin posteriorly. Possibly some of the larvae are merely faded specimens of one of the other types. This is a realistic prospect with the two Naga specimens from the Sulu Sea (USNM 337337) which are faded; however, the remaining specimens are not faded. The absence of a second Pt spine, late formation of Scl spine three, predominance of specimens of this type from the western Indian Ocean, and the well-developed pigment on the base of the caudal rays all suggest that this larval type does indeed constitute a distinct population.

eraseziS.eavralih-hpromoiseacaraPfosepytehtgnitaitnereffidsretcarahctnemgiP.3elbaT yrevsihcihw,)7.giF,2elbaT(eanilispAdeifitnedinUotoslarefeR.mmtseraenehtotdednuor otralimis oiseacaraP .tnenimorpotsuoucipsnocniyrevsetacidnielbairaV.ih-hprom

)egnarezis(epytlavraL ABC DF ()Fig.2)()Fig.3()Fig.5()Fig.3-4(Fig.6 S4izerange(mm)33-131-143-134-14-2 PIGMENTCHARACTER Doorsal-finbasenonsyoensye Aonal-finbasensyseyseyseye Lsowerjaw yoensyseyoen Uerostyle vtariablptrominenptrominenptrominenprominen B;asecaudalrays0x-100x-3;fle0x-2;fle0x-2;fle1-3;fle (Numberofusually>1inusually1;0usually1;0usually1;0usually2;0 melanophores)bothflexandinpostflexafter7mmafter9mmafter7.5mm postflex LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 709 ;egalna,a;tneipicni,i;degamad,d.mmnistnemerusaeM.eavralAepytih-hprom e n i I p s e / .)selacsLL57tuoba(selacssahnemicepsmm59.31ehtylnodna,senipsOSevahsnemicepsehtfoenoN.sunata,)A(;esablarotcepta,)P( n . . . i O 51 41 31 p PO ss a r o t i 15 1d1 151 15 fs c e PO a , , , , , , , I I I I I I i n I I I I I I I fn I 941 I5 I6 I3 I0 Ii5 Ii41 Al a s 1 1 1 1 1 1 1 , r i , , , , , , Iii20 fn o Vii31 X Xd X8 X8 X8 X9 X8 I8 Dl t s g 0 0 6 4 2 . . . . . n 2 20 20 0V 11 10 e Pl ln t p g 8 s . n 55 e Dp lh A ( d 0 8 0 0 3 7 3 3 h ...... o t 3dd0 1dd0 2d8 1ddII 1dd0 0d8 1d0 1d6 p B2 e dh P ( oiseacaraP d 6 2 2 3 3 7 7 7 h ...... o t 31 2d6 24 24 13 14 15 17 p By e d) t u g o 8 9 8 8 4 5 fosretcarahccitsiremdnacirtemohpromdetceleS.4elbaT 3 5 6 ...... n n 07 05 05 09 08 00 0d6 03 05 e Sy l) e t i e b 2 0 9 1 6 6 3 7 7 r ...... m 15 13 04 15 05 01 06 09 04 a Ot i dh t a g 5 9 7 9 6 6 1 9 1 e ...... n 36 20 27 20 14 19 18 17 29 e Ht lr a s t r g o 4 7 7 7 7 1 0 0 ...... n d 38 24 24 16 19 13 29 23 e e lh r Pd a t n g 5 5 8 3 3 5 7 8 a ...... n e 38 33 37 23 22 14 21 27 r e lh Pl N N O I O I X X E N E L O h I F L 5 t ydo F T X 9.3 51. 6 2 6 1 03. 4 g S E E 6. 4. 6. 1 2 n R L O 8 . . e Bl P lh 14 ~dd6 F 77 77 51 58 P 39 52 60 710 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Most specimens of type A have much less urostyle pigment than is usual in Paracaesio morph-hi larvae, whereas a few have the usual pigment pattern. Too few specimens are available to evaluate this variation in pigment: it could represent geographical variation, individual variation, or even the presence of more than one species within this type. Ad- ditional material of type A is required.

Paracaesio morph-hi type B 49 specimens, 2.7-13.0 mm Figure 3A, Tables 3, 5

Material Examined. —East China Sea, AMS I.31824-007, I.31824-008, I.31832-003, I.31979-005, I.31980-002, I.31987-002, I.31998-003, I.32041-003, I.32042-003, I.32042-004, I.32042-005; MUFS 11293, 11294, 11295; NSMT-PL 37, 38: Ryukyu Isls., AMS I.31544-001, I.31803-045: Banda Sea, LACM 44496-106: eastern Indian Ocean off Sumatra, ZMUC DANA 3867-I-IV, DANA 3881-IV: western Indian Ocean, AMS I.33008-006, I.33152-004.

Diagnosis. —MORPHOLOGY - morph-hi; second spine on dorsal ridge of Pt absent; third spine on Scl forms early (from 5.5 mm). PIGMENT. - No dorsal pigment on trunk or tail. No melanophore at tip of lower jaw. Melanophore present at base of A fin posteriorly. Remarks. —This larval type is very similar to Paracaesio morph-hi D (see Remarks under type D, below). In type B the third Scl spine is first present at 5.5 mm, and is possibly consistently present from that size. However, we lack specimens between 5.9 and 8.5 mm, so some specimens larger than 5.5 mm may lack the third spine.

Paracaesio morph-hi type D 38 specimens, 3.0-13.3 mm Figures 3B, 3C, 4, Tables 3, 7

Material Examined. —Sagami Bay, Japan, CBM-ZF 5970: East China Sea, AMS I.31824-010, I.31979-006, I.32013-004, I.32042-007, I.32087-008: western Pacific near Palau, AMS I.31527- 004, I.31808-001: Bismarck Sea, AMS I.19753-068: Makassa Strait, ZMUC DANA 3793-II, DANA 3796-III: western Coral Sea, AMS I.30510-001, I.32534-002: western Tasman Sea, AMS I.26129- 009, I.26131-013, I.26161-003, I.28817-001: eastern Indian Ocean off Sumatra, ZMUC DANA 3863- I-III, DANA 3867-I-IV, DANA 3875-II, DANA 3896-I,II: western Indian Ocean, AMS I.33053-010, I.33123-004, I.33135-005, I.33180-005; RUSI 48812.

Diagnosis. —MORPHOLOGY - morph-hi; second spine on dorsal ridge of Pt absent; third spine on Scl forms late (about 7 mm). PIGMENT. - No dorsal pigment on trunk or tail. Melanophore present at tip of lower jaw. Melanophore present at base of A fin posteriorly. Remarks. —Larval type D is very similar to type B, differing only in a slightly later formation of the third Scl spine (in type D first present at 6.9 mm, and consistently present from 8.4 mm, whereas in type B first present at 5.5 mm), and in the presence of a melanophore on the tip of the lower jaw. The two differences, albeit slight, between larval types B and D suggests they represent different species. Both larval types have similar, wide distributions in the western Pacific and Indian Oceans, the only obvious difference being the absence of type B in collections from off LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 711

Figure 3. Larvae of Paracaesio morph-hi type B and D. Type D has a melanophore on the lower jaw, and type B lacks this pigment (see text for other differences). Scale bar is 1 mm. A. Type B, 2.65 mm from the western Indian Ocean (AMS I.33008-006). B. Type D, 4.15 mm from the western Indian Ocean (AMS I.33123-004). C. Type D, 4.50 mm from the western Indian Ocean

(AMS I.33053-010). P2 ray 1 is broken. 712 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

the east coast of Australia. The known distributions of P. sordida (absent off Australia) and P. xanthura roughly correspond to those of Paracaesio morph-hi types B and D, respectively (Allen, 1985), but distributions of Paracaesio are too poorly known to base identifications of larvae solely upon them.

Paracaesio morph-hi types C and F 41 specimens, 4.3-23.5 mm Figures 5, 6, Tables 3, 6

Material Examined. —Sagami Bay, Japan, CBM-ZF 5969, 5970: East China Sea, AMS I.31533- 001, I.31534-001, I.31824-009, I.31824-011, I.31832-004, I.31979-007, I.31999-002, I.32042-006, I.32085-002, I.32087-007, I.32089-009, I.32090-007, I.32090-008; ZMUC DANA 3722-IV, DANA 3723-V, DANA 3727-III; NSMT-PL 39, 40; MUFS 11296, 11297, 11298: western Pacific near Palau, AMS I.31548-002, I.31788-001, I.31807-001, I.31809-001: Banda Sea, LACM 44471-6, 44496- 57; ZMA 121.533: eastern Indian Ocean off Sumatra, ZMUC DANA 3875-II, DANA 3869-VIII.

Diagnosis. —MORPHOLOGY - morph-hi; second spine on dorsal ridge of Pt only rarely present, if present, forming at about 7.5 mm; third spine on Scl forms late (6.5 mm). PIGMENT. - Pigment present posteriorly at base of D fin, initially a single melanophore on each side, later increasing to a cluster of melanophores, and in the largest larva spreading ventrally over the side as a weak bar. Melanophore on lower jaw either present (type C) or absent (type F). Melanophore present at base of A fin posteriorly. Remarks. —Types C and F may represent different species, but we could find no other differences that correlated with the presence or absence of the melanophore at the lower jaw, and therefore, we describe them together. We had no specimens of type C larger than 11 mm, suggesting the possibility that type C looses the lower jaw melanophore at about that size, becoming indistinguishable from type F. The two larval types have similar distributions in the western Pacific, but type F was collected only in the NW Pacific, whereas type C was also taken in the Indonesian Archipelago, including waters of the extreme eastern Indian Ocean off Sumatra. We identified no preflexion specimens of either type C or F, suggesting the possibility that the dorsal pigment in these types does not form until the time of flexion. Larval Aphareus and Pristipomoides (eteline lutjanid genera) have pigment on the tail dorsally similar to that of Paracaesio morph-hi C and F, and in these genera the dorsal pigment does not form until late in the preflexion stage or during the flexion stage (Leis and Lee, 1994), providing a precedent for this suggestion. More larval material, and a clearer understanding of the systematics and distributions of Paracaesio spp with about 70 LL scales are needed to resolve the identities of types C and F. LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 713

Figure 4. Larvae of Paracaesio morph-hi D. Internal structure in fin spines not shown. A. 5.90 mm from the Bismark Sea (AMS I.19753-068). P2 ray 1 is broken. B. 10.8 mm from the eastern Indian Ocean off Sumatra (ZMUC Dana 3875-II). Both Dsp2 and P2 ray 1 are broken. Proximal pigment on Dsp2 is in the chevron groove, distal pigment is on the membranous sheath surrounding the spine (sheath is missing on the most distal portion of the spine). Note pigment on soft rays of caudal and dorsal fins. Specimen is largely covered by developing scales, but only those along LL are shown. 714 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997 ;egalna,a;tneipicni,i;degamad,d.mmnistnemerusaeM.eavralBepytih-hprom e n i p s e / n i O 21 p POI ss e l a LO c ss a r o t fs c e PL a n Al a s I r I i I 00000 I 00000 I 00010 I 00010 Ia0010 Ia0010 Ia0020 I 00000 Va0031 Vai031 fnifni Vii031 Vii031 Vii031 o Vii031 Dl t s g 2 3 2 2 3 2 2 3 6 7 7 8 7 9 9 8 ...... n 2 0I 0I 0I 0I 0I 0I 0I 0I 0Va005. 0Va0031 0I 0I 0Vii031 0Vii031 0I 0Vii031 e Pl ln t p g 7 7 6 7 6 7 8 2 1 4 1 s ...... n 04 05 06 06 09 07 05 29 26 20 20 e Dp lh A ( .senipsOSevahsnemicepsehtfoenoN.sunata,)A(;esablarotcepta,)P( d 4 6 6 5 6 5 6 6 7 9 8 8 9 9 9 9 9 9 0 0 h ...... o t 01 02 09 02 0d1 09 04 08 0d8 0d5 05 0ddI 0d6 00 0d6 03 02 0ddVii031 1ddI 1ddI p B2 e oiseacaraP dh P ( d 6 7 7 7 8. 7 7 8 0 1 1 h 1 1 2 2. 2 2. 3 4 ...... o t 02 09 01 08 05 08 07 01 15 11 17 10 16 12 11 15 12 10 12 p By e fosretcarahccitsiremdnacirtemohpromdetceleS.5elbaT d) t u g o 1 2 1 1 2 2 2 3 3 3 4 4 4 4 4 3 4 ...... n n 02 06 07 08 07 09 06 04 01 05 00 04 05 0d 2 08 01 00 e Sy l) e t i e b 3 3 3 3 3 3 3 5 5 5 5 5 5 5 5 5 6 6 r ...... m 05 01 08 08 01 03 02 00 07 09 00 00 05 00 00 0d 5 09 04 a Ot i dh t a g 6 7 8 8 9 8 9 9 2 3 3 4 3 4 3 4 4 5 5 6 e ...... n 05 08 05 05 0dd4 05 09 09 14 16 10 13 10 16 10 1dd3 16 11 12 14 e Ht lr a s t r g o 8 9 9 9 9 8 0 0 2 3 3 3 3 4 4. 3 4 6 5 6 ...... n d 07 08 02 02 04 00 15 19 16 18 14 12 16 12 17 15 14 10 14 16 e e lh r Pd a t n g 1 2 2 2 3 2 2 3 6 8 8 8 8 9 0 0 0 0 2 a ...... n e 11 13 18 12 19 15 10 17 16 14 12 14 15 13 27 24 22 22 22 r e N lh Pl O I X E htgne L ydo F 5 86. 27. 57. 67. 08. 38. 19. 93. 47. 57. 28. 58. 09. 00. 60. 60. 14. 4 16. E 6 5 R . . Bl lh P 20 20 26 26 20 26 24 26 39 30 30 35 30 39 4d0 41 41 48 48 44 LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 715 e n i p s e / n i 11 O 31 51 41 51 71 p POI ss e l 7 7 6 a 70 LO ~71 ~91 ~91 c ss a r o t 105. 105. 1i61 1i81 11 10 1i71 19 12 fs c e PL a , , , , , , , , , , , , , ,I , , ,I , , , 1 , , I , I , , I I I , I I I I I I I I n , I I I I I I I I Ii041 I Ii05. Ii05. I I I Ii041 I I I I I I I I I I8041 Ii021 Ii041 I605. I8041 Ii031 Ii031 Ii041 I8061 I9041 I8041 I0 I7 I4 I6 I7 I7 I6 I7 Ii041 Idi81 I7 Al Ii041 a 1 , , , s I 1 I 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 I , , , , r , , , , , , , I I i , , , , , , , , , , , , , I I I Vii021 fnifni o I X X X X X X X X X X X X9 X8 X9 X8 X8 X8 X8 X8 X8 X8 X8 X9 X8 I8 I9 I9 I9 I8 I9 I8 I0 I9 I8 I9 V6 V8 Dl V7 t s g 2 4 1 2 9 5 9 2 0 2 9 5 2 6 1 7 4 1 7 3 ...... n 2 10 10 17 11 09 10 01 11 20 20 01 20 10 20 11 20 20 10 20 11 e Pl ln t p g 4 5 4 s . . . n 37 63 37 e Dp lh A ( d 9 4 0 3 0 3 0 5 9 4 0 9 1 1 1 3 0 6 2 6 1 3 6 1 7 2 3 4 h ...... o t 0ddI 1d0 1dd0 1d7 12 1d5 1d4 1d5 0d1 1d5 1dd1 1d0 1dd0 20 1d6 2d5 10 2d2 1d2 2d2 1dd1 2d1 2dd0 1d3 2dd0 3d6 1d3 3dd0 p B2 e dh P ( d 3 7 4 7 4 7 4 8 4 8 4 5 5 9 4 1 5 0 6 1. 5 2 0. 6 3 7 h 8 1 ...... o t 19 14 14 19 18 12 11 10 16 10 16 23 19 23 13 36 19 30 16 30 17 36 36 17 34 33 13 44 p By e d) t u g o 5 5 4 6 4 5 4 5 4 6 4 7 5 9 5 9 5 0 5 5 0 5 1 0 5 0 ...... n n 09 02 02 05 06 02 05 05 04 05 06 09 08 06 07 04 08 17 02 02 15 02 11 16 05 17 e Sy l) e t i e b 5 7 6 7 6 7 6 8 6 8 6 0 6 2 7 2 6 3 7 3 6 2 7 5 3 5 8 7 r ...... m 01 09 03 00 06 07 06 08 06 05 08 17 01 11 03 16 03 10 03 1d 0 03 10 07 11 1d 0 15 09 19 a Ot i dh t a g 6 1 7 1 6 9 6 2 5 1 8 9 7 1 8 2 8 5 9 7 8 4 9 0 7 1 0 7 e ...... n 19 25 17 29 17 15 17 22 17 21 11 21 18 30 10 30 18 34 12 34 19 36 14 45 32 41 21 40 e Ht lr a s t r g o 6 0 9 1 6. 1 7 2 5 1 7 8 7 1 9 1 7 5 8 5 8 2 9 7 6 2 1 5 ...... n d 16 25 13 21 11 21 19 20 15 20 19 20 16 35 17 38 11 35 13 31 16 37 15 33 31 40 25 44 e e .deunitnoC.5elbaT lh r Pd a t n g 2 6 5 7 7 3 8 3 1 3 9 3 6 4 2 4 3 4 0 4 1 6 2 5 8 7 3 a ...... n e N 26 25 20 29 24 25 25 25 30 25 31 27 46 21 43 21 55 25 56 26 53 26 61 53 58 25 74 r e O lh I Pl X E N L O htgne I F 42.1 78.1 6 ydo T X 5 8 06. 6 1 3 56. 2 07. 98. 5 4 97. 95. 8 29. 5 85. 51. 87. 5 87. 2 88. 5 9.2 S E 4 2. 3. 6. 4. 5. 7. 5. 9. 0. 2 5. 7 L O . . . Bl lh 40 54 F 40 52 4d7 53 40 52 48 53 45 86 49 84 41 81 51 98 50 90 51 93 52 13 90 12 56 11 P 716 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 5. Larvae of Paracaesio morph-hi type C. Type C has a melanophore on the lower jaw, and type F (Fig. 6) lacks this pigment (no other differences were found). Internal structure in fin spines not shown. Scale bar is 1 mm. A. 4.8 mm type C from the East China Sea (AMS I.31824-009). P2 ray 1 is broken. No internal structure in fin spines. B. 7.0 mm type C from Sagami Bay, Japan

(CBM-ZF 5969). P2 ray 1 is broken, but the length is estimated from a 7.3 mm specimen. Weak internal structure present in Dsp2 & 3, P2sp and Asp2. C. 9.9 mm type C from the East China Sea (AMS I.32087-007). Specimen has a large number of developing scales, those shown are to indicate size and spacing, not extent of coverage. Internal structure present in all fin spines. P2 ray 1 is broken, and a portion of the membranous sheath of Dsp2 is damaged. Note pigment on caudal rays. LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 717

Figure 6. Larvae of Paracaesio morph-hi type F. Type C (Fig. 5) has a melanophore on the lower jaw, and type F lacks this pigment (no other differences were found). Blank area posterior to edge of hypural plate indicates where scales cover caudal rays. Internal structure in fin spines not shown. Scale bar is 1 mm. A. 13.9 mm type F from the East China Sea (MUFS 11296). Specimen is fully scaled, but only the LL pores are shown. Internal structure present in all fin spines. Dsp2 and P2 ray 1 are broken. Note pigment on the caudal rays. B. 23.5 mm type F from the Philippine Sea (AMS I.31788-001). Myomeres are not shown because they are obscured by the full set of scales; of these, only the LL pores are shown, but the blank area at the base of the caudal rays indicates coverage by scales. Internal structure is present in all fin spines. The tip of P2 ray 1 is damaged. Note pigment on soft rays of D, A, and C fins. 718 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997 tuohtiwnemiceps)*(.mmnistnemerusaeM.eavralFepytdnaCepytih-hprom snemicepsehtfoenoN.sunata,)A(;esablarotcepta,)P(;tneidicni,i;egalna,a;degamad,d;detardyhernemiceps,+;)Fepyt=(wajrewolnoerohponalem e n i I p s e / n i O 31 41 51 p PO ss e l a LO c ss a r o t fs c e PL a , , , ,I , , , , I n I I I I I I i041 I I 4041 I i0d1 I 4041 I605. I i041 I i041 I605. Al a s , , , 1 , 1 1 1 , , r i , , V i i031 fnifni X X X X o V i i020 Vii05. V i i021 X X I8 I7 I6 I7 X9 X8 I9 I9 V 6i031 Dl t s g 2 3 1 1 3 1 3 0 ...... n 2 19 18 18 10 11 19 10 11 e Pl ln t p g 4 1 1 0 2 s . . . . . n 34 3dI 3d 8,III 44 38 e Dp lh A ( d .ylevitcepser,8dna2evahhcihw,tsegralowtehttpecxesenipsOSevah 9 9 0 2 9 1 1 9 1 2 2 4 1 h ...... o t oiseacaraP 0ddII 0ddII 1ddII 13 00 1d5 1d6 09 1d2 10 1d6 1d7 14 p B2 e dh P ( d 4 3 3 5 4 5 6 3 6 7 6 8 6 h ...... o t 13 17 11 11 19 16 18 17 14 17 15 15 15 p By fosretcarahccitsiremdnacirtemohpromdetceleS.6elbaT e d) t u g o 4 4 4 5 4 5 5 4 5 6 5 5 5 ...... n n 04 02 01 03 04 07 00 09 02 09 05 02 02 e Sy l) e t i e b 6 5 6 6 5 6 7 6 7 8 7 6 7 r ...... m 07 07 07 05 08 01 07 08 07 03 02 01 04 a Ot i dh t a g 4 5 6 7 6 8 9 6 9 0 9 9 8 e ...... n 12 18 16 17 19 14 13 11 10 20 19 16 10 e Ht lr a s t r g o 5 5 7 9 7 9 9 7 9 9 9 0 9 ...... n d 15 16 13 17 15 18 11 14 12 10 10 25 19 e e lh r Pd a t n g 9 0 0 3 3 3 4 3 6 6 4 7 6 a ...... n e 19 27 22 20 24 29 24 28 26 25 20 26 21 r e N lh Pl O I X NOIXEL E h L 5 23.4 41.5 04.5 34.5 t ydo F 2 62. 08. 36. 41. 42. 83. 26. 17. g E . n R 4 e *4 46 *4 48 43 59 F *3 54 54 *5 *9 Bl 53 53 lh P LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 719 e n i I p s e / n i 11 13 14 O 41 51 61 41 61 61 71 91 71 p 11 PO ss e l 6 7 a 75. LO ~91 ~91 c ss a r o t 0 105. 105. 105. 105. 1d5. 1 061 1i71 1i71 1i81 17 1d 5.0 1i71 1i5. 10 1271 13 1172 1179 fs c 1 0d1 e PL a , , , , , , , , , , , , , , , , , , , , , , , , , , ,I I I I I I I I I I I I I I I I I I I I I I I I I n I I I I I I I I I I I I I I I I I I I I I I I I I I I I i041 I 5051 I505. I 6051 I905. I 5051 I1 I 8041 I+ I1 I3 I4 I d0d1 I4 I4 I2 I4 I7 I7 I7 I7 I7 I7 I6 I6 I6 I7 Al a s 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1, 1 1 1 1 1 , r i , , , , , , , , , , , , , , , , , , , , , , , fnifni o X X9 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 I9 Dl t s g 2 3 4 9 7 4 9 9 1 0 3 2 5 5 7 0 7 2 9 8 ...... n 2 11 10 10 10 10 10 10 10 20 20 20 20 20 20 20 30 30 30 30 40 e Pl ln t 4 p g 5 4 6 1 6 3 9 5 . s ...... n 4 36 57 40 62 52 50 72 8d0 e Dp 16 lh A ( d 1 3 5 4 7 3 5 4 7 5 6 7 1 7 1 5 5 0 9 1 6 4 3 7 h ...... o . t 1d1 17 1d2 1dd0 1d2 1dd0 12 10 1dd0 1dd0 17 10 2d0 12 2d3 2d3 24 3d6 20 3d7 3d0 3d3 4d6 63 p B2 e dh P ( d 3 7 8 0. 1 9 2 0 3. 9 3 4 4 5 7 0 h 2. 3 7 5 7 4 7 6 3 ...... o . t 15 16 15 29 21 10 28 29 20 15 27 21 26 20 24 30 3dd7 38 39 35 35 47 49 42 76 p By e d) t u g o 5 6 6 6 6 7 8 8 7 7 8 7 8 9 8 1 0 0 1 3 4 1 ...... n n 06 07 05 04 00 00 04 09 03 02 08 08 01 00 04 12 10 12 15 12 18 23 e Sy l) e t i e b 6 7 7 7 8 8 8 8 0 9 1 1 9 1 1 2 2 3 4 4 6 5 8 4 r ...... m 07 02 04 0d 5 08 05 08 07 14 04 15 18 08 11 15 1d 1 15 10 15 17 13 19 14 23 a Ot i dh t a g 8 2 1 5 4 2 5 4 7 7 7 9 9 7 1 5 7 8 8 1 2 3 0 2 8 e ...... n 10 24 21 21 27 25 27 29 29 25 2dd9 20 22 24 32 35 30 35 32 40 45 44 59 58 73 e Ht lr a s t r g o 6 2 0 3 3 3 3 2 5 5 6 7 8 8 7 2 2 5 7 7 0 5 6 0 6 ...... n d 13 21 25 27 26 22 24 25 29 27 27 22 20 28 25 33 32 38 38 33 45 44 44 55 79 e e lh .deunitnoC.6elbaT r Pd a t n 8 g 2 7 6 2 0 1 4 2 6 5 5 9 9 7 8 5 3 0 2 5 6 4 9 0 . a ...... n e 0 N 29 21 29 39 33 39 39 36 30 32 35 32 30 35 37 48 45 59 55 54 50 60 68 78 r e 12 lh O Pl I X E L +01.1 98.31 54.32 h F 57dddd d dd 01, +5.7~ddddd 72.5 14.6 15.6 55.6 23.7 44.7 37.7 3 t 92.1 83.1 33.3 29.4 ydo T 7 40. 68. 11. 94. 99. 57. 31. 57. 00. 88. 1.1 g S 4 n O . e 45 Bl 57 lh P *2 50 69 *5 60 *0 *8 63 69 *3 *2 *3 88 80 98 90 1 ddddd d15 d dd 01, 15 13 18 *0 15 *5 720 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997 ,)P(;egalna,a;tneipicni,i;degamad,d.mmnistnemerusaeM.eavralDepytih-hprom e n i I p s e / n i O 31 21 p PO ss e l a LO c ss a r o t i fs c e PL a , , , , I I ifn n Ii031 I I I I Ii041 Ii041 Ii041 Ii041 Al a s I I 1 1 I I 1 1 , , r I I i , , Vii031 Vii0 5. I a0020 I a0020 I a0021 I a0021 fn o Vii05. V i0031 X X Vii041 VI Vii041 X8 X8 I9 I9 Dl t s g 0 9 9 9 7 7 0 4 4 5 4 8 3 2 7 0 ...... n 2 1I 0I 0I 0I 0V 0 Vi0031 1I 0V 11 0V 10 0V 10 11 0 Va0031 1I e Pl ln t p g 1 4 7 6 3 1 s ...... n 32 37 10 15 26 31 e Dp lh A ( d 9 0 1 0 7 8 8 6 8 3 7 3 7 2 1 0 7 0 h ...... o t 0d2 1d6 14 16 00 0d4 0d6 0d8 0 ddVa0021 1d0 00 1d6 0d0 1d6 1d0 1ddIII 00 10 p B2 e .senipsOSevahsnemicepsehtfoenoN.sunata,)A(;esablarotcepta dh oiseacaraP P ( d 2 2 2 2 0 2 1 9 9 7 9 6 9 6 5 4 9 2 h ...... o t 11 17 10 19 16 18 15 05 04 15 07 11 04 15 18 10 08 18 p By e d) fosretcarahccitsiremdnacirtemohpromdetceleS.7elbaT t u g 4 4 4 4 3 4 4 o 3 3 5 3 5 3 6 5 4 2 4 ...... n n 00 03 02 05 08 08 09 04 06 02 07 08 05 09 06 08 01 07 e Sy l) e t i e b 6 6 5 5 4 5 5 3 5 8 4 8 3 7 7 6 4 6 r ...... m 07 04 04 03 06 01 05 03 05 08 00 07 06 00 04 08 07 08 a Ot i dh t a g 6 5 5 5 2 4 4 1 3 1 9 9 1 1 9 7 1 7 e ...... n 15 10 17 14 19 13 10 17 14 20 00 10 19 28 10 14 16 12 e Ht lr a s t r g o 7 6 4 5 2 4 3 2 2 0 1 1 0 1 8 7 1 6 ...... n d 16 13 11 12 14 18 17 14 15 20 10 28 10 21 12 11 13 10 e e lh r Pd a t n g 2 1 8 9 7 0 8 5 7 6 4 7 4 7 5 1 4 2 a ...... n e 29 22 19 14 19 22 13 11 15 23 13 21 15 23 29 21 18 23 r e lh N Pl O I X NOIXEL E htg L ydo F 78. 76. 54. 71. 51. 10. 68. 47. 44. 3 04. 6 92. 3 11. 85. 3 84. E 5 4 4. 0 n R . . . e F 45 49 49 44 45 40 35 39 39 55 35 51 38 58 55 49 32 41 Bl lh P LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 721 e n i I p s e / n . i O 61 51 51 41 p PO ss e l 7 7 7 7 a LO ~81 ~81 ~71 ~81 c ss a r o t i 10 12 10 10 1i71 1i71 1i71 1061 1071 105. 105. 1071 105. 1051 1051 fs c e PL a , , , , , , , , , , , , , , , , , , , , I I I I I I I I I I I I I I I I I I I I ifn n I I I I I I I I I I I I I I I I I I I I I7 I7 I6 I6 I dd71 I6 I6 I6 I5 I3 I3 I3 I1 I4 I3 I 9061 I0 I 9061 Ii05 I d041 Al a s 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 r i , , , , , , , , , , , , , , , , , , , , fn o X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 Dl t s g 6 7 7 2 3 2 1 2 2 0 2 9 7 7 7 6 2 ...... n 2 20 20 20 20 20 20 20 20 20 20 20 10 10 10 10 10 10 e Pl ln t p g 8 1 5 8 s . . . . n 63 63 51 57 e Dp lh A ( d 0 8 6 4 3 2 2 2 1 8 2 8 8 7 8 5 5 3 3 h ...... o t 3d5 2d6 2d1 2dd0 2d6 2d9 2d2 2d2 2d0 1d5 2d0 11 1d9 12 1dd0 1d3 12 1dd0 1d8 p B2 e dh P ( d 5 6 0 1 7 9 9 6 7 7 4 6 3 3 1 1 0 9 7 8 h ...... o t 32 38 35 36 2d5 29 23 25 23 28 27 20 24 26 27 26 24 16 11 16 p By e d) t u g 0 9 0 9 8 8 8 o 9 8 7 7 8 7 7 7 7 7 6 6 6 ...... n n 15 00 15 07 05 04 02 05 06 03 00 09 00 02 09 09 04 02 05 01 e Sy l) e t i e b 3 3 3 2 1 1 1 0 1 0 9 0 9 9 9 7 9 8 8 8 r ...... m 18 15 10 17 11 11 15 15 13 19 06 16 05 05 02 00 00 00 05 06 a Ot i dh t a g 6 6 5 5 1 2 0 2 1 9 8 8 6 5 6 5 4 1 2 2 e ...... n 35 31 36 34 30 38 35 32 31 20 26 25 27 29 27 27 28 24 22 25 e Ht lr a s t r g o 9 5 6 1 2 0 0 0 0 9 7 8 7 3 4 5 4 2 1 2 ...... n d 38 30 37 34 39 31 34 32 37 27 21 27 29 27 21 20 29 29 26 22 e e lh r .deunitnoC.7elbaT Pd a t n g 3 1 1 3 2 4 2 3 1 9 8 8 1 4 3 2 1 0 8 8 a ...... n e 58 55 54 45 48 49 44 42 40 38 34 31 44 39 37 32 35 39 25 25 N r e lh O Pl I X E L htg F 61.1 18.0 51.0 ydo T 91. 38. 47. 56. 84. 04. 10. 15. 84. 42. 29. 98. 35. 72. 09. 76. 3 S 5 n O . e 14 15 15 96 84 81 84 86 84 87 70 71 70 65 68 61 69 55 58 Bl 53 lh P 722 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Unidentified Apsilinae 19 specimens, 2.1-6.2 mm Figure 7, Tables 2, 8

Material examined. —central Pacific, Tuamotu Islands, AMS I.29035-002, I.29075-003, I.29078- 004: western Coral Sea, AMS I.29996-006, I.30609-009, I.30625-012: western Tasman Sea, AMS I.26073-001, I.26073-002, I.26080-003, I.26110-012, I.26120-014, I.26124-006, I.26129-010, I.28816-001, I.31392-004.

Diagnosis. —MORPHOLOGY - Similar to Paracaesio morph-hi; body moderately deep (20-37% BL); head spines moderate in number and size; PAsp reaches maximum of 11% BL; Dsp2 very slender and very long (to 80% BL), much longer than other Dsp (Dsp3 7- 22% BL); in no specimen is Dsp2 enclosed by distal sheath with small, fleshy “flag” at its

tip; P2sp of moderate length (7-28.6% BL), but P2 ray 1 nearly as long as Dsp2 (182- 275% P2sp). PIGMENT. - No pigment on midbrain dorsally; usually no melanophore posteriorly at A base, but if present, very small; if present, ventral pigment on tail limited to 1-2 melanophores posteriorly on caudal peduncle after middle of preflexion stage (3.7 mm); no melanophore at cleithral symphysis; small external melanophores on urostyle from flexion. Description. —With the following exceptions, morphology and pigmentation (Table 2) are the same as that described for Paracaesio morph-hi. Some head spines form slightly later, the sequence of (and size at) formation of the first of these spines on the head bones is as follows: interopercle (4.2 mm), supracleithrum (4.0 mm), and posttemporal (5.0 mm). The least developed specimen has yet to develop fin spines unlike the least developed Paracaesio morph-hi larvae which had Dsp2 and 3, however the latter is larger. The Unidentified Apsilinae larvae have no pigment that does not also occur in other Paracaesio morph-hi larvae, but they lack some, most notably the midbrain melanophores, and have smaller and fewer melanophores in most other locations. Remarks. —There are two apparent differences between Paracaesio morph-hi larvae and the Unidentified Apsilinae larvae concerning ornamentation of the very elongate fin elements that are probably artefacts. The absence of a sheath with a pigmented “flag” on Dsp2 in the relatively small number of Unidentified Apsiline larvae available may be an artefact of damage rather than a real difference between them and the other Paracaesio morph-hi larval types. A very high percentage of Paracaesio morph-hi larvae lacks this sheath, apparently due to damage, although it occurs in at least some specimens of every

larval type. Similarly, a single Unidentified Apsilinae larva has a pigmented “flag” on P2 ray 1 (Fig. 7B). This is not present in any of the Paracaesio morph-hi larvae we examined, but Okiyama (1991) illustrated it in what appears to be a Paracaesio morph-hi type

F larva. The elongate ray of the P2 fin is broken off in the majority of larvae examined. A number of specimens with the ray unbroken have tiny bits of pigment (e.g., Fig. 7C), perhaps representing the remnants of a pigmented “flag”. The pigmented “flag” is possibly present in all morph-hi larvae, but so fragile that it is almost never found intact in preserved material. Otherwise, the Unidentified Apsilinae larvae are essentially morphologically identical to the Paracaesio morph-hi larvae. It is most likely they represent a species of Paracaesio, perhaps P. pedleyi McCulloch and Waite (1916) originally described from the Tasman Sea-Coral Sea area, if it is distinct from P. xanthura (Anderson, et al. 1992; Anderson, LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 723

Figure 7. Larvae of the Unidentified Apsilinae. Scale bar is 1 mm. A. 2.8 mm from off Takapoto Atoll in the Tuamoto Islands (AMS I.29078-004). B. 3.2 mm from off Osprey Atoll in the western Coral Sea (AMS I.30609-009). Right eye illustrated in place of missing left eye. C. 5.1 mm from the Western Tasman Sea (AMS I.26073-002). Tip of P2 ray 1 is damaged, only a tiny amount of pigment remains. 724 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997 ;tneipicni,i;degamad,d;nekorbylbissop,*.mmnistnemerusaeM.eavraleanilispAdeifitnedinUfosretcarahccitsiremdnacirtemohpromdetceleS.8elbaT e n i I p s e / n i O 20 p PO ss a r o t i 141 fs c e PO a , , , , , , , I i n I I I I I941 Ii41 I fn I531 I641 I841 I631 I1 Al a s 1 1 1 1 , 1 1 , , , , , r I I I i , I0000 I0000 I0000 I0020 I0010 Ia030 Ia020 I005. fn X o X X X X X I8 X8 I9 I9 I9 I9 I9 Dl t s g 3 3 3 3 8 8 6 6 7 5 4 5 4 2 0 3 3 1 7 ...... n 2 11 11 11 11 0Va030 0V 0V 0V 0I 0I 0I 0I 0I 0I0000 0I0000 0I0000 17 11 10 e Pl ln t 3 p g 0 8 7 8 5 4 4 9 6 5 2 6 9 . s ...... n 3 40 33 21 21 18 13 17 00 11 00 00 09 29 e Dp *5 lh .senipsOSroselacsevahsnemicepsehtfoenoN.sunata,)A(;esab1Pta,)P(;egalna,a A ( d 4 3 2 3 7 9 6 8 7 6 6 5 6 5 3 3 2 0 2 3 h ...... o t 1d3 1d4 14 14 00 08 02 0d1 05 0d6 07 07 09 00 01 05 0--a0000 1d0 15 19 p B2 e dh P ( d 7 8 5 5 0 2 9 9 8 7 7 6 7 5 5 5 4 5 6 9 h ...... o t 13 19 10 10 18 12 04 07 02 08 09 00 03 00 04 09 09 18 16 19 p By e d) t u g o 5 6 5 6 5 4 3 3 2 1 2 3 2 2 1 6 4 8 ...... n n 00 01 06 05 07 05 02 07 00 09 08 08 05 05 01 02 00 07 e Sy l) e t i e b 7 7 6 6 5 5 4 5 4 3 3 3 3 3 2 6 5 7 r ...... m 07 04 02 07 00 09 04 08 07 08 03 00 00 02 05 05 01 01 a Ot i dh t a g 9 0 8 8 9 4 2 3 9 7 9 8 9 8 7 7 5 8 4 4 e ...... n 18 26 14 13 04 11 14 13 00 08 07 0dd6 09 00 04 0dd6 01 14 18 24 e Ht lr a s t r g o 1 0 8 8 4 4 2 3 0 9 1 9 9 0 8 8 9 5 3 ...... n d 28 28 14 11 19 15 10 10 13 05 17 08 04 10 01 01 19 19 23 e e lh r Pd a t n g 5 8 2 3 8 9 6 6 3 1 2 3 2 3 2 2 9 4 0 3 a ...... n e 21 25 24 27 16 16 10 19 13 15 15 18 19 10 13 17 0-6 24 24 38 r e lh Pl N N O I O I X X E N E L O htg I F L ydo F T X 1 72. 4 9 21. 10. 27. 17. 12. 11. 00. 09. 7 0 6 0 7 7 5 8 S E E 5. 0. 7. 8 8 6 4 0 7. 2. 1. ne R L O . . . . . P lh Bl P 56 56 54 46 F 48 43 38 32 39 30 39 21 25 20 21 27 28 44 47 61 LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 725

1994). However, because the largest specimen is only 6.2 mm and lacks scales, we cannot be certain this series is not Parapristipomoides squamimaxillaris, which has an adult distribution in many respects similar to that of the Unidentified Apsilinae larvae and whose larvae are undescribed. Further, no Paracaesio species is known to have an adult distribution similar to that of these unidentified apsiline larvae (the specific status of the Paracaesio species reported from French Polynesia by Randall (1985) is uncertain, although it is a member of the P. xanthura complex (Anderson, 1994)), but too much weight should not be placed on this in view of our poor knowledge of Paracaesio distributions. From outside the western Coral Sea-Tasman Sea area (i.e., the east coast of Australia), we have only five specimens, 2.1-3.0 mm BL (the most developed of which has Dsp III), of this larval type. They are from off Takapoto, an atoll in the Tuamotu Archipelago, French Polynesia. There are some slight pigment differences between the Takapoto specimens and the Australian material. The five Takapoto specimens have an internal melanophore dorsally on the hindbrain and a melanophore at the tip of the lower jaw (Fig. 7A). Of the eight Australian specimens of equivalent developmental state, only four have the hindbrain melanophore, three have the lower jaw melanophore (although all specimens larger than 3.7 mm have it), and three of the eight have both melanophores. The material is too limited to determine if this pigment variation is due to small sample size, to geo- graphic variation or to the putative series containing more than one species. The Unidentified Apsilinae larvae are very lightly pigmented, and because only relatively small (<7 mm) larvae are available, it is possible they merely represent smaller, less heavily pigmented specimens of one of the Paracaesio morph-hi types of larvae. How- ever, the size range of all of these other types partially overlap that of the Unidentified Apsilinae larvae. Further, the unique distribution of these larvae (South Pacific as far south as Sydney, Australia), suggests that at the very least, they are a distinct geographical variant of one of the other species. It seems likely they constitute a distinct species.

Paracaesio morph-lo Table 2, Figures 8-13

Diagnosis. —MORPHOLOGY (values in [ ] indicate postflexion individuals).- body deep (28-46[38-46]% BL); head spines moderate in size; PAsp reaching maximum of 15% BL; Dsp2 robust and not remarkably long (10-31[14-29]% BL), only slightly longer than

Dsp3 if at all (Dsp3 5-23[14-23]% BL); Dsp2 without distal sheath or “flag” at tip; P2sp of moderate length (11-28[18-27]% BL); P2 ray 1 slightly shorter to somewhat longer than P2sp (90-171[97-127]% P2sp); scales beginning to form at about 7 mm, fully formed at about 9 mm; about 50 LL scales. PIGMENT. - Melanophore at cleithral symphysis present from about 2.6 mm to at least 4 mm, variably present until 5.0-6.6 mm, depending on species; no pigment at urostyle; dorsal pigment on trunk or tail, if present, not externally associated with posterior end of dorsal-fin base; pigment on dorsal fin, if present, limited to chevron groove and associated membrane of Dsp2 until about 9 mm when it may be present distally on membranes of other spines and rays; P2 pigment sparse, and if present, limited to mid to distal portions of rays 1-2, sometimes spreading to other rays after 5 mm. Description. —MORPHOLOGY - Body compressed and moderate to deep (28-46% BL). Youngest larvae somewhat more elongate, and in postflexion larvae BD 38-46% BL. Body depth changes gradually by development of a steeper profile to forehead and con- 726 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

comitant deepening of trunk myomeres to give hunch-back appearance, with maximum

relative depth at about 9-10 mm. The body is deeper at P2 base than at anus. Gut mor-

phology and changes in position of P2 fin same as in morph-hi. Head large and compressed. The snout length less than eye width. Snout initially pointed, but as head profile steepens, snout becomes blunt. Mouth large, somewhat oblique and reaches at least anterior margin of eye, but not to mid eye. Small, conical, teeth first form anteriorly on premaxillae at about time Dsp2 is forming. Dentary teeth form soon after. A small canine tooth anteriorly in premaxilla forms at 8-12 mm. No large canines form. Nasal pit bridges over forming two nostrils between 7 and 9 mm. Spination on head moderately developed. All head spines smooth, and except for a single, small serra (or small spine) on supraorbital ridge (FS) in one species, and serrations on the lachrymal in another, no ridges have serrations (serrations on preopercular outer border seen late in larval stage formed by large increase in number, but no increase in size, of spines). Least developed specimen (2.55 mm) has only spine at preopercular angle and one on upper limb of inner border of preoperculum. Sequence of (and size at) formation of first spine on other bones is interopercle (3.5 mm), supracleithrum (3.5 mm), posttemporal (3.5 mm), subopercle (10 mm). Spine at angle of preopercle largest on head, reaching 4-15% BL. It gradually becomes engulfed by border of preopercle, but is prominent in largest specimen. Inner border of preopercle has up to 4 small spines, and all present by 4.5-6.6 mm, depending on species. Spines on opercle, interopercle, and outer border of preopercle as in morph-hi. Interopercular spination forms initially as small spine just dorsal to and approximately parallel to spine at preopercular angle. Up to four additional small spines added to border of interopercle from 12 mm. Up to six small spines on subopercle from about 14 mm. A single opercular spine appears at about 3.5 mm.

A small spine forms on dorsal postcleithrum just dorsal P1 base at about 4.5-5 mm at about time flexion complete, and persists in largest specimen. Supracleithrum develops its first spine at about 3.5 mm and adds two more small spines. Small posttemporal spine appears at about 3.5 mm and up to three spines present on dorsal ridge from about 9 mm. On ventral ridge of posttemporal a small, single spine forms early in postflexion stage, with a second present in some individuals from 9 mm. Supraorbital ridge develops at about 3.5-4.0 mm and is present in largest individual. In most larvae, ridge remains smooth, except for slightly hooked posterior edge from about 9 mm. In one species, however, a single, small supraorbital spine forms over mid eye at about 4.7 mm. Low, smooth frontal ridge dorsal and slightly anterior to supraorbital ridge forms at about 7 mm. A low, smooth pterotic ridge forms at about 5 mm approximately in line with supraorbital ridge. Ridges on supratemporal-intertemporal as in morph-hi. Maxilla develops at anterior end a spine-like acute angle. At about 9 mm, lachrymal begins to project over posterior end of maxilla and to cover the maxilla when mouth closed. The border of the lachrymal not smooth, but without spines or serrations except in largest specimens (>28 mm) where distinct serrations present in at least one type.

First fin element to form is P2 spine at about 2.9 mm. Shortly thereafter Dsp2 forms.

Most of D spines and P2 rays form before D soft rays, elements of A fin and primary caudal rays begin to form. However, latter all present by completion of notochord flexion. Flexion begins at about 4.0 mm and is complete at about 4.8-5.4 mm. Except for Dsp1, D spines form from anterior to posterior. Ultimate D spine initially a soft ray, but remains so over very short range of BL, and transforms into spine before flexion complete or very shortly thereafter. Asp3 forms initially as soft ray early in flexion stage and LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 727

transforms into spine during flexion or very shortly thereafter. P1 rays last to form, and all

present at about time scales form (sometime between 7 and 9 mm). Dsp2, P2 spine and P2

rays 1 and perhaps 2 become very elongate: Dsp2, 5.8-31.2[14-29]% BL; P2sp, 10.7-

28.3[18-27]% BL; P2 ray 1, 90-171[97-127]% P2sp. Dsp2 slightly longer than P2sp until

about 5.5 mm, after which they are about equal until about 12 mm, thereafter P2 longer.

Maximum relative spine length reached between 5 and 9 mm. P2 ray 1 very slightly

shorter to somewhat longer than P2sp. A spines not particularly elongate. All fin spines chevron-shaped in cross-section, and concave (trailing) surface contains tissue that is

sometimes pigmented. On leading edge of P2 spine and anterior D spines are two ridges basally and single ridge distally. Basal split ridges first obvious at 5-5.5 mm, and extend

about 10-20 % (P2sp) or 20-30 % (Dsp1) of the length of the spine. No fin spines orna-

mented externally, but distinctive internal structure develops during flexion in D and P2 spines (Figs. 1C,D). Internal structure first in D spines during flexion, becomes strong at

about 6 mm, and present in all dorsal-fin spines by about 9 mm. Structure in P2 strong at about 9 mm. Internal structure forms at about 5.7 mm in A spines becoming strong at about 12 mm. Gill rakers 18-22 (5-6+1+12-15) between 9 and 10 mm, 19-24 (5-7+1+13-16) between 11.5 and 12.6 mm, 22-23 (6+1+15-16) between 14.0 and 17.8 mm, 25 (8+1+16) at 19.1 mm, 26 (8+1+17) at 20.3 mm, and 25 (7+1+17) at 28.6 -29.4 mm. Adults of this morph have 26-32 (7-12+1+15-19) gill rakers (Table 1), so by about 14 mm adult complements of gill rakers on lower limb are attained. In contrast, on upper limb, adult complements not attained until after 30 mm. Scales form relatively early. Incipient scales present from about 7 mm. A gap in the series makes it impossible to be precise about when scales all present, but full complements of scales present by 9 mm. Scales on maxilla may form much later. Smallest specimen with scales on maxilla is 28.6 mm, however not all species have scaled maxilla and few of our larvae are identified to species, so it is not possible to be certain about size at which scales form on maxilla. PIGMENT. - Pigment generally light. Pigment cap present dorsally on gas bladder and gut. Some preflexion specimens have a small melanophore on preanal fin fold at anus. Pigment dorsally on midbrain varies with species, but present in most specimens, and occurs at a very small size. Dorsal melanophores increase in number and spread to cover midbrain by about 9 mm. No melanophores forebrain until about 9 to 12 mm (depending on species), but by 14 mm they cover it. At tip of lower jaw a melanophore always present until 7.2 mm, and variably present until 11.7 mm. An internal melanophore appears at base of opercular spine at about 7 mm. Melanophore at cleithral symphysis varies among species: one species has a melanophore at cleithral symphysis between 2.9 and 6.6 mm, and the other has one variably present between 4.0 and 5.2 mm. Larger specimens lack cleithral melanophore. Initially a series of up to eight small melanophores along ventral midline of tail. These decrease in number to at most two melanophores by completion of flexion, one at posterior end of anal fin and one near posterior end of caudal peduncle. Both become internal, with anterior one usually disappearing by 5.6 mm, posterior melanophore more likely to be present: however, both present in some specimens up to 19 mm. No pigment at urostyle. Pigment on dorsal surface of trunk and tail limited and varies among taxa. Compared to other lutjanids, pigment on fins very limited until late in larval development. On D fin, pigment initially absent, and may remain so, but specimens as small as 728 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 8. Morph-hi larva of Paracaesio gonzalesi or kusakarii 28.6 mm from the Pacific Ocean off the north coast of Irian Jaya (AMS I.34957-001). Strong internal structure in the spines of the fins is not shown. Myomeres are obscured by scales and are not shown. Specimen is fully scaled, but scales are shown only on the maxilla (diagrammatically) and along the lateral line (pores only). Blank area posterior to edge of hypural plate represents scales covering caudal rays. Note pigment on D and A fins. Scale bar is 1 mm. 3.5 mm have small amount of pigment associated with distal portion of Dsp2 and membrane. From 9 mm this may spread to membranes of other spines, and by 12-14 mm, there may be distal band of pigment along membrane of spiny portion of fin. In larger specimens, fin membrane may have scattering of melanophores proximal to band. On A fin, largest specimens with distal pigment on membrane of anterior portion of fin (Fig.

7). Pigment on P1 absent, and on caudal fin, it is limited to a small melanophore initially on anlage and later on base of fin (either dorsally or ventrally), but even this may be

absent. Most specimens lack pigment on P2 fin, but it may be present in larvae between 4 and 12 mm in form of one to five small melanophores on central portions of ray one or two, or in very large larvae on ray 5. Remarks. —In larvae of Paracaesio morph-lo, the numbers of gill rakers on the lower limb meet or exceed the values found in adult Lipocheilus by the time the larvae are 11.5 mm. The different types of Paracaesio morph-lo differ by minor morphological and pigment characters (Table 9). At least three species seem to be represented among our material.

Paracaesio gonzalesi Fourmanoir & Rivaton, 1979 and/or kusakarii Abe, 1960 3 specimens, 28.6-29.4 mm Figure 8, Tables 9, 10

Material Examined. - Pacific Ocean off north coast of Irian Jaya, ZMUC DANA 3768-XIV; AMS I.34957-001.

Diagnosis. —MORPHOLOGY - morph-lo; serrate lachrymal; 50 LL scales; scaled maxilla. BD 39% BL, Dsp2 14-16% BL, P2sp 19-20% BL. PIGMENT. - Six dorso-lateral bars (or saddles) of diffuse melanophores positioned at midbrain, nape, bases Dsp2-3, bases LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 729

fosepyttnereffidehtgnitarapesnilufesusretcarahC.9elbaT oiseacaraP .eavralol-hprom

epytlavraL ADC- gonzalesi/kusakarii S0izerange(mm) 39-249-28-2 CHARACTER Smallspineonsupraorbital roidge nsyoen Soerrationsonlachrymalnonsye Socaledmaxilla n?sye G2ill-rakersat8-9mmBL280-21? Boroaddorso-lateralbarsn?sye Dorsalpigmentoncaudalvariableto6.6nsoonlybar peduncle mm,thenabsent Dsorsalpigmentonmidbrainy5evariabletoyes mm,thenyes V,entralpigmentontailyesuntil5-9mmuosuallyyesn occasionallylarger C6leithralmelanophoreyesfrom3-6.variableuntil5no mm,thenabsentmm,thenabsent

Dsp8-9, base D ray 5 and caudal peduncle (longest bars extend to just ventral of LL);

membrane of D fin with distal band of pigment from Dsp1 to middle of soft rays; P2 usually with a small area of pigment on ray 5. Remarks. –Relative to smaller morph-lo specimens, these specimens have reduced head spination, with the frontal ridge absent, and the other ridges small, PAsp about 5% BL and PI spines absent. The number of PO, IO and SO spines is high, but all are small, approaching serrations. Obviously, these fish are in transition from the larval to the juvenile stage. Lachrymal serrations have not been reported in Paracaesio of any stage previ- ously, and may represent a late larval or juvenile character. This larval type has four distinct serrae on the lachrymal. The other Paracaesio morph-lo larvae have only the weakest projections along the lachrymal, and very few, if any, of these would be considered serrations or spines. Similarly, none of the other Paracaesio morph-lo larvae have scales on the maxilla or pigmented bars on the body. It is not clear at what size these characters form in P. gonzalesi and/or kusakarii, and it may be that even the largest specimens of the other Paracaesio morph-lo types (20 mm) are simply too small to have them. The distributions of both P. gonzalesi and P. kusakarii overlap broadly (Allen, 1985), and the larvae were caught within the known or expected ranges of both.

Paracaesio morph-lo type A-C 46 specimens, 2.6-20.3 mm Figures 9-11, Tables 9, 11

Material examined. - East China Sea, AMS I.31540-006, I.31788-002, I.31824-012, I.31824-013, I.31832-005, I.31832-006, I.31978-005, I.31980-003, I.31982-002, I.31999-003, I.31999-004, I.32017-001, I.32018-001, I.32041-004, I.32041-005, I.32063-002, I.32084-008, I.32088-003, I.32090-009; ZMUC DANA 3722-IV,V, DANA 3727-III; NSMT-PL 41; MUFS 11299, 11300: Pacific Ocean off north coast of Irian Jaya, AMS I.34956-001; ZMUC DANA 3775-II: Molucca Sea, LACM 36065-68: Banda Sea, ZMA 121.534, 121.535. 730 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 9. Larvae of Paracaesio morph-lo type A-C from the East China Sea. No internal structure in fin spines. Scale bar is 1 mm. A. Type B, 2.55 mm (AMS I.31824-013). B. Type A, 2.95 mm (AMS I.31832-005). C. Type A, 3.60 mm (AMS I.31824-014).

Diagnosis. —MORPHOLOGY – morph-lo; no spines on FS; lachrymal without distinct

serrations; about 50 LL scales; BD 28-46% BL; Dsp2 6-29% BL; P2sp 4-27% BL; PAsp 5-13.5% BL. Pigment - Dorsal pigment on caudal peduncle present in many specimens <6.6 mm BL. Some pigment always present dorsally on midbrain, and in preflexion larvae, some dorsolateral pigment at mid-hindbrain border moves ventrally to behind otic LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 731 ,a;tneipicni,i;degamad,d.mmnistnemerusaeM.eavral e n i p s e n i IOS p ss e n i 236 235.5 p 245 PO ss e l a 54 54 55.6 LO/O c ss a r o t 10 10 10 fs c e PL a n fni I6 I7 I6 Al a s 1 1 1 r i , , , fni o X8,II X8,II X8,II Dl iselaznog.P t s g 6. 5. 8. n 2 50 50 50 e Pl ln t p g 4 1 3 s . . . n 44 41 45 e ro Dp lh iirakasukoiseacaraP A ( 1 0 9 d . . . h o t 0 0 0 p B2 15 11 17 e dh P ( 2 1 3 d . . . h o t 1 1 1 p By 14 18 12 e d) fosretcarahccitsiremdnacirtemohpromdetceleS.01elbaT t u g o 8 9 6 . . . n n 24 24 28 e Sy l) e t i e b 4 4 2 r . . . m 38 36 32 a Ot i dh .sunata,)A(;esablarotcepta,)P(;egalna t 7 9 a g . . 7 e . n 0 0 91 e Ht 16 19 lr a s t r 3 3 7 g o . . . n d 0 0 0 e e 18 11 14 lh r Pd a t n 4 9 1 g . . . a n e 6 6 7 r e 17 17 13 lh Pl N O I X E L htgne F 0 07.8 04.9 ydo T 6 S . O 8 Bl 22 lh P 28 21 732 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 10. Larvae of Paracaesio morph-lo type A-C from the East China Sea. Scale bar is 1 mm. A. Type A, 4.50 mm (AMS I.31824-014). No internal structure in fin spines. B. Type B, 5.62 mm

(AMS I.31999-003). Internal structure in Dsp2 and P2 not shown. LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 733

Figure 11. Larvae of Paracaesio morph-lo type A-C. All are type B. Internal structure present in all spines, but not shown. Scale bar is 1 mm. A. 9.68 mm from the Molucca Sea (LACM 36065- 068). Specimen fully scaled anteriorly, but most scales are missing leaving only scale pockets. The only scales illustrated are those remaining in place on LL. B. 12.6 mm from the Molucca Sea (LACM 36065-068). Specimen is fully scaled, but only LL pores are shown. The Sb spine shown is from the right side; in the illustrated specimen, it is absent on the left side. C. 19.1 mm from the East China Sea (AMS I.32088-003). Specimen is fully scaled, but only scales along LL are shown; blank area posterior to edge of hypural plate indicates scales covering caudal rays. 734 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997 ;egalna,a;tneipicni,i;degamad,d.mmnistnemerusaeM.eavralC-Aepytwol-hprom e n i p s e n i IOS p ss e n i p PO ss e l a LO/O c ss a r o t fs c e PL a , , , I n I I I Ii0410 Ii0410 Ii0410 Al a s I 1 1 1 I I I , , , r I i Ia00200 fnifni o Vi00310 Vi00410 VI00310 VI00310 X X X Vi00410 I9 I8 I9 Dl t s g 7 6 7 5 5 1 2 1 9 1 8. 2 1 8 ...... n 2 0I 0I 0I 0Va00310 0Va00310 0ia00200 0aa00200 0ia00200 01 11 01 0I 0aa00200 0I e Pl ln t p g 8 7 8 5 6 1 2 0 1 8 s ...... n 02 04 00 01 09 19 15 1dII 07 07 e Dp lh A ( d 1 1 9 . 7 6 6 6 4 4 2 0 5 6 5 6 4 2 h ...... o t 01 15 17 03 04 0-7 0-7 0-6 17 13 1d4 12 07 0-1 0--aa00000 0--aa00000 0 --a-00000 16 p B2 e dh oiseacaraP P ( d 5 5 5 1 1 9 9 9 7 9 6 5 8 8 7 7 7 6 h ...... o t 15 11 18 18 11 01 07 03 16 12 19 12 07 07 01 01 06 10 p By e d) fosretcarahccitsiremdnacirtemohpromdetceleS.11elbaT t u g o 4 4 4 3 3 2 2 2 5 6 5 4 3 2 2 2 1 4 ...... n n 00 03 06 04 07 02 02 00 04 01 09 05 06 04 03 04 06 01 e Sy l) e t i e b 5 5 5 4 5 4 4 4 6 7 6 5 4 4 3 3 3 5 r ...... m 03 04 06 09 05 06 06 06 08 07 02 00 00 04 04 02 05 04 a Ot i dh t a g .sunata,)A(;esablarotcepta,)P( 5 5 5 2 3 9. 1. 9. 8 1 8 6 1 9. 8. 7. 8. 5 e ...... n 17 17 16 15 10 03 12 00 18 28 16 19 13 00 06 00 04 18 e Ht lr a s t r g o 6 6 4 3 1 8 0 7 7 1 7 ...... n d 14 15 16 17 16 17 24 14 10 10 15 e e lh r Pd a t n g 2 0 2 7 8 5 4 4 3 8 4 2 4 4 1 1 2 2 a ...... n e 24 22 25 16 19 1-9 1-1 1-3 26 24 28 20 10 1-4 1-3 1-4 1-1 21 r e lh Pl N O I X NOIXEL E htgne L ydo F 21. 50. 16. 74. 23. 50. 79. 05. 73. 13. 05. 59. 4 4 55. 5 40. E 6. 9 6 5 R . . . F 44 47 37 37 35 30 36 21 45 54 40 40 28 23 25 25 20 41 P lh Bl LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 735 e n i p s e n i IOS 15.0 p ss e n i 145.2 125.2 112 121 15. 110 110 110 135.5 133 111 p PO ss e l a 55.6 55.5 54 53 43 4910 LO/O ~4 ~1 ~0 ~0 ~0 ~910 c ss a r o t 10 10 10 194 10 19 105 194 174 194~5.0 18 1d910 154 194 1i610 10610 10610 10510 10510 10510 10610 10510 10410 fs c e PL a ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II ,II , ,II ,II ,II I n I I6 I6 I6 I6 I6 I7 I5 I6 I6 I6 I6 I6 I6 I6 I3 I1 I1 I2 I2 I1 I1 I2 I90410 I80510 I0 I70410 I60410 I60410 Al a s 1 1 1 1 1 1, 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 , r i , , , , , , , , , , , , , , , , , , , , , , , , fnifni o X X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 X8 I8 Dl t s g 0 6 8. 5. 9 5 6 7 3. 5. 6 3 4 9. 4 3. 5 5. 3 4. 0 1 1. 1 9 ...... n 2 40 30 30 20 20 20 20 20 20 20 20 20 20 10 10 10 10 10 10 10 10 11 10 10 00 e Pl ln t p g 1 2 7 7 6 7 6 4 3 7 7 7 3 3 4 3 3 0 s ...... n 40 35 20 26 21 21 20 29 24 14 1d01, 17 19 13 1d01, 11 13 13 e Dp lh A ( d 9 1 4 2 0 4 0 9 0 9 7 5 4 4 7 1 3 0 9 1 6 7 7 9 7 6 7 6 h ...... o t 63 68 5d6 5dd0 5d3 40 40 35 43 3d6 3d0 30 37 35 2d3 24 20 2d5 16 2d4 15 1d2 16 11 11 1d7 16 12 p B2 e dh P ( d 8 1 8 0 7 2 8 6 8 5 4 1 1 0 2 6 6 5 6 6 1 2 2 3 2 0 0 9 h ...... o t 72 78 64 65 52 50 48 46 45 44 41 40 49 45 37 23 20 22 26 20 25 24 26 24 22 28 21 17 p By e d) t u g 9 6 7 4 2 1 1 1 1 0 9 9 o 8 8 6 6 7 7 5 7 5 6 6 6 5 5 6 5 ...... n n 11 13 17 10 18 10 10 18 10 17 06 05 00 00 07 05 02 02 05 01 05 02 00 05 06 09 00 06 e Sy l) e t i e b 4 3 2 1 0 7 7 5 5 5 4 4 2 2 0 0 9 9 8 8 8 8 7 7 6 7 7 7 r ...... m 22 21 21 26 23 15 14 10 16 11 15 18 15 19 12 18 07 02 08 03 06 05 08 00 09 04 03 03 a Ot i dh t a g 5 7 5 0 4 1 6 7 5 2 0 1 4 5 6 6 7 4 3 4 2 6 2 1 9 0 1 0 e ...... n 75 61 64 66 51 55 48 42 49 41 46 45 37 37 25 20 26 22 25 27 21 27 29 25 18 28 20 23 e Ht lr a s t r g o 1 4 5 6 5 9 7 2 3 6 8 8 6 5 7 5 7 4 4 4 2 2 0 2 1 9 0 0 ...... n d 77 67 60 55 54 40 45 40 45 30 34 39 35 33 29 23 21 23 23 21 25 20 25 28 29 15 25 23 e e lh r .deunitnoC.11elbaT Pd a t n 2 3 7 g . . . 4 1 1 9 6 5 6 7 5 3 3 8 5 6 2 2 4 9 1 9 7 7 8 7 5 a ...... n 1 0 0 e 83 87 70 68 67 64 56 51 57 58 53 33 37 34 31 34 34 24 37 29 23 29 29 29 22 r e 13 12 10 lh N Pl O I X E L htgne F 03.0 60.9 77.7 95.4 00.4 06.2 53.2 86.1 94.1 50.0 ydo T 37. 86. 90. 10. 42. 06. 03. 21. 90. 50. 17. 46. 26. 63. 50. 10. 49. 1 S 8 O . 29 19 11 12 19 10 18 12 18 14 93 95 90 95 79 63 65 69 65 61 58 54 52 57 58 52 41 P 43 lh Bl 736 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 12. Larvae of Paracaesio morph-lo type D. Scale bar is 1 mm. Internal structure in fin spines not shown in B. A. 4.03 mm (identification tentative - note lack of spine on FS ridge) from the western Coral Sea (AMS I.32520-001). No internal fin-spine structure. B. 4.67 mm from the

East China Sea (AMS I.31824-015). Internal structure in fin spines: Dsp2 weak; P2sp very weak. Dsp10 and Asp3 transforming from ray to spine.

capsule, remaining there until at least 10 mm. Melanophore at cleithral symphysis be-

tween 2.9 and 6.6 mm. Most specimens lack pigment on P2; if present, only 1-2 melanophores. Pigment usually present ventrally on tail until about 20 mm. No dorsolateral bars on body. Description. —Pigment dorsally on midbrain in smallest specimens. This consists of 4-5 melanophores at mid-hindbrain border, at least one of which on or near dorsal midline, and the others of which more lateral in position. Dorsal melanophores increase in number and spread to cover midbrain. Most of more lateral melanophores quickly move ventrally to laterally along mid-hindbrain border behind otic capsule. Initially a number of melanophores along the ventral midline of the tail, but these decrease in number so that by end of flexion, only two melanophores remain, one at posterior end of A fin and one near posterior end of caudal peduncle. Both become internal, with anterior one usually disappearing by 5.6 mm, whereas posterior melanophore more likely to be present in some specimens up to 19 mm. Included here are larvae with some variations in pigment, particularly pigment of the dorsal surfaces of the trunk and tail. Some of smallest larvae (<2.7 mm) have one or two LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 737

small melanophores mid-dorsally on trunk. These melanophores appear to move internally and become lost to view or disappear. On dorsal midline of peduncle, some larvae up to 4.5 mm have a melanophore just posterior to base of dorsal fin, and some larvae up to 6.6 mm have a melanophore on penultimate myomere (these are type A, Figs. 9B,C, 10A). Again, both melanophores appear to move internally before becoming lost to view or disappear. A few specimens lack posterior melanophore on ventral midline of caudal peduncle (these are type C, not illustrated). Type B (Figs. 9A, 10B, 11A-C) lacks dorsal pigment on peduncle, but has ventral melanophore lacking in type C. Remarks. —The series has a gap between 7.2 mm and 9.0 mm, and the 7.2 mm specimen is the only one between 6.6 and 9.0 mm. This limits the precision of our description in some areas, particularly scale formation. There are only 5 specimens larger than 12.6 mm, but they are spread between 12.6 and 20.3 mm (Table 11). Therefore, the description is reasonably complete. A question remains as to how many species might be included in this series. Morphologically, there is no indication of more than one species, but pigment is variable, and this could indicate that more than one species is involved. The largest type A is only 6.6 mm, leading to the supposition that the distinguishing dorsal melanophore is lost at larger sizes. Distributions of adults of Paracaesio morph-lo are too incompletely known to be helpful in establishing identifications.

Paracaesio morph-lo type D 7 specimens, 4.0-9.2 mm Figures 12-13, Tables 9, 12

Material Examined. - East China Sea, AMS I.31824-015; NSMT-PL 42: western Pacific near Palau, AMS I.31800-001: western Coral Sea, AMS I.32520-001: Banda Sea, LACM 44496-107.

Diagnosis. —MORPHOLOGY - morph-lo; small spine on middle of FS; lachrymal with-

out distinct serrations; about 50 LL scales; BD 31-46% BL; Dsp2 10-31% BL; P2sp 11- 28% BL; PAsp 4-15% BL. PIGMENT. - No dorsal pigment on peduncle. Pigment on dorsal surface of midbrain absent in some specimens up to 4.9 mm. No dorsolateral pigment at mid-hindbrain border before scales form. Cleithral symphysis melanophore variably

present between 4.0 and 5.2 mm and absent thereafter. Pigment present on P2. Ventral pigment on tail disappears sometime between 5.2 and 9.2 mm. Remarks. —The distinguishing feature of this larval type is the single, small spine on FS. In the largest specimen, this spine seems to be reduced in size, and it may be absent in larger larvae. Morph-lo type D apparently differs from morph-lo type A-C in several pigment features (Diagnosis, Table 9), but this could be an artifact of low numbers of specimens. The smallest specimen lacks the FS spine (Fig. 12A), and it is thus only tentatively identified, but it was in the same sample as the 5.23 mm (Fig. 13A) specimen and has the morph-lo type D pigment features. The series of only seven specimens is very incomplete with no specimens larger than 9.2 mm and a gap between 5.23 and 9.2 mm. This limits the detail that can be provided, but general development seems to differ little, if at all, from that of P. morph-lo type A-C. The largest morph-lo type D larva (9.16 mm) has 5+1+12 = 18 gill rakers, and this is lower by 2-4 rakers than the count in 9-10 mm larvae of type A-C larvae. The type D 738 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

Figure 13. Larvae of Paracaesio morph-lo type D. Scale bar is 1 mm. Internal structure in fin spines not shown. A. 5.23 mm from the western Coral Sea (AMS I.32520-001). Weak internal

structure in Dsp2. P2 pigment is that of the right fin. B. 9.16 mm from the Banda Sea (LACM 44496-107). Strong internal structure in fin spines: Dsp1-6 strong, 7-10 weak; Asp1-3 weak; P2sp moderate. Specimen is scaled, but only the lateral-line scales are shown.

count is within the range of values recorded for Lipocheilus carnolabrum adults (Table 1), and this raises the possibility that morph-lo type D is Lipocheilus, and not Paracaesio. However, based on other apsiline larvae, a 9 mm specimen would be expected to have 2 to 8 lower-limb rakers fewer than the adult complement. If this applies to morph-lo type D, then the adult complement would exceed that of L. carnolabrum. In addition, two of the morph-lo type D larvae are from the western Coral Sea (16° 28' S), outside the known LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 739 ;egalna,a;tneipicni,i;degamad,d.mmnistnemerusaeM.eavralDepytwol-hprom e n i I p s e / n i O 51 31 31 91 p PO ss a r o t 15. fs c e PO .selacsLL94sahhcihwtsegralehttpecxe,selacsrosenipsOSevahsnemicepsehtfoenoN.sunata,)A(;esablarotcepta,)P( a ,I ,I , ,I ,I ,I I n I I I I I I fni I95. I55. I45. I6 I741 I841 Al a s 1 1 1 1 1 1 , r i , , , , , fni o X X8 X8 X8 X8 X8 I8 Dl t s g 3 1 1 5 4 3 1 ...... n 2 10 10 11 20 0Vaa30 10 10 e Pl ln t p g 6 3 4 4 3 3 s ...... n 15 12 20 03 12 14 e Dp lh A ( d 8 4 3 4 9 4 7 h ...... o t 12 13 1d0 35 00 12 14 p B2 e dh P ( d 3 7 9 2 2 9 1 h ...... o t 20 18 16 40 15 18 27 p By e oiseacaraP d) t u g o 5 4 4 9 3 5 6 ...... n n 00 07 00 00 07 01 04 e Sy l) fosretcarahccitsiremdnacirtemohpromdetceleS.21elbaT e t i e b 7 8 6 4 4 7 7 r ...... m 02 09 07 15 06 02 00 a Ot i dh t a g 2 0 9 1 3 9 1 e ...... n 24 20 16 40 18 10 25 e Ht lr a s t r g o 0 1 8 6 4 9 1 ...... n d 26 21 11 33 17 17 25 e e lh r Pd a t n g 5 5 5 2 8 5 5 a ...... n e 26 20 26 58 11 22 26 r e lh Pl N N OIXELFTSO O I X N E OIX htg L ydo F 02. 19. 9 6 3 76. 32. E E 6. 1. 0 n R L . e P lh P 56 Bl 44 43 98 F 43 41 57 740 BULLETIN OF MARINE SCIENCE, 61(3): 697–742, 1997

range of L. carnolabrum. However, the adult lower-limb gill-raker complement is estab- lished by 9 mm in some other lutjanid larvae, e.g., some Pristipomoides spp. (Leis and Lee, 1994). Further, the distribution of L. carnolabrum is poorly known, and could well extend to the western Coral Sea: the nearest published records are in New Britain (Kailola, 1987) and north-western Australia (Allen and Swainston, 1988). Finally, there are so few differences in head spination among Paracaesio species larvae that it is possible the supraorbital ornamentation is generically important. So, although it seems unlikely based on present evidence, there is thus a possibility that the larvae identified here as Paracaesio morph-lo type D are in fact L. carnolabrum. Larger larvae of this taxon are required to resolve this.

DISCUSSION

The two morphs of Paracaesio larvae are very different (Table 2). Paracaesio morph- hi larvae are of similar body shape to eteline lutjanid larvae (Leis and Lee, 1994), and have only moderately developed spination on the head. In contrast, Paracaesio morph-lo larvae have much deeper bodies, and have relatively larger, and earlier-forming head spines. The scales of morph-hi larvae are in the incipient stage over a long period, and begin to form at a size when morph-lo larvae already have a full complement of scales. However, the most obvious difference between the two morphs is in fin-spine morphol-

ogy. Morph-hi larvae have extremely long, pigmented Dsp2 and P2 ray 1, and the other fin spines are slender (Figs. 1C,D). In contrast, the fin spines of morph-lo larvae are robust, relatively short and have little or no pigment. Internal structure in the fin spines is weaker in morph-hi than in morph-lo. Larvae of Paracaesio morph-hi are unlikely to be confused with any other lutjanids as

long as either Dsp2 or P2 ray 1 is wholly or largely intact. These elements are frequently broken, but even in this case, it is often possible to ascertain the distinctive round cross- sectional shape of distal portions of Dsp2. Otherwise, Paracaesio morph-hi larvae are likely to be confused with some etelines, especially so as some species of each have similar pigment patterns (Leis and Lee, 1994). The fin-ray counts of each are diagnostic, but until the fins form, if the elongate elements are broken, confusion is likely to occur. The deep-bodied larvae of Paracaesio morph-lo are superficially similar to a number of lutjanine larvae with smooth fin spines. Once the fins form, the diagnostic counts of DX,10 and AIII,8 will identify the apsiline larvae. Also, lutjanines typically do not have pigment on the dorsal surface of the midbrain at as small a size as do apsilines. Most Paracaesio morph-lo larvae lack serrations on the supraorbital ridge (except type D which has one small serra), whereas these are present in most, if not all, known lutjanines at least in the postflexion stage. Finally, in lutjanines, the second and subsequent spines of the outer, upper preopercular border do not form until well after flexion is complete (generally, >8 mm) whereas in apsilines they are added from early in the postflexion stage. The very large differences between the two larval morphs of Paracaesio are consistent with the suggestion of Anderson, et al. (1992), based on adult characteristics, that there are two monophyletic groups within Paracaesio. The very elongate, morphologically distinct, fin elements of the morph-hi species are unique among the lutjanids, and are certainly derived within the family, thus providing good evidence of monophyly of this group. The same may prove true of the rather long time (implied by size differences) LEIS ET AL.: PARACAESIO LARVAL DEVELOPMENT 741 between initiation of scale development and its completion. At present, it is more difficult to say if any of the character states of the morph-lo larvae are derived. The very deep body shape of morph-lo larvae, possibly the deepest in the family, is conceivably derived, but more work is needed to corroborate this. This leaves open the possibility that Paracaesio morph-lo is a paraphyletic group or is based on plesiomorphic characters. On the other hand, with the possible exception of the distal band of pigment in D and A fins of late larvae (Figs. 6B, 8 and 11C), no evidence from larvae suggests that Paracaesio (sensu Anderson et al., 1992) is a monophyletic entity. At the same time, we have no clear evidence from larvae that Paracaesio is not monophyletic. Nor is there any evidence available from study of adult morphology (Anderson et al., 1992; Anderson, 1994), although W.D. Anderson, Jr. (pers. comm.) believes that Paracaesio is monophyletic. The unique (for lutjanids) combination of fin-ray counts (DX,10; AIII,8) provides the only evidence of monophyly of the Apsilinae as presently understood, but this must be regarded as very tentative until we have a better understanding of the sister group(s) of the Lutjanidae. We are aware of no evidence from larvae indicating that the traditional subfamilies Etelinae and Apsilinae are distinct, monophyletic entities. It must be kept in mind that we have not seen larvae of all nominal apsiline genera, but given the differences among larvae within Paracaesio, it seems unlikely that larval morphology will provide evidence of apsiline monophyly.

ACKNOWLEDGMENTS

This study could not have proceeded without the generous cooperation of many people who made available the collections 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. E. Beckley, E. Bertlesen, K. E. Hartel, Y. Iwatsuki, G. D. Johnson, J.-I. Kojima, M. Miya, K. Matsuura, G. E. McGowan, A. J. Miskiewicz, J. Nielsen, M. Okiyama, T. Ozawa, J.R. Paxton, B. Ranchod, W. J. Richards, R. H. Rosenblatt, T. Saruwatari, P. Schalk, J. A. Seigel, L. D. TerMorshuizen and H. J. Walker. S. E. Reader and T. Trnski ably assisted in the lab. W. D. Anderson, Jr., shared his unsurpassed knowledge of apsiline lutjanids. He and C. C. Baldwin constructively criticized the manuscript. M. McGrouther made sure our material was registered in good time. A. Hay registered many of those specimens as did B. Ewart who also made Fig. 1 and assisted with editorial chores. T. Goh assisted with data entry and production of the typescript. This project was supported by Australian Research Council grant A19031159, and by the Austra- lian Museum. The collection of many specimens from the western Coral Sea was supported by Australian Marine Science and Technology grant 86/0873 to J.H. Choat and J.M.L. Our very great thanks to all.

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DATE ACCEPTED: March 29, 1996.

ADDRESSES: Fish Section, Division of Vertebrate Zoology, Centre for Evolutionary Research, .The Australian Museum, 6 College Street, Sydney, NSW 2000, Australia. PRESENT A DDRESS: (S.B.) Royal Botanic Gardens, Mrs. Macquaries Road, Sydney, NSW 2000, Australia.