Larvae of Five Fish Families with Patfern 10 Of

BULLETIN OF MARINE SCIENCE, 60(1): 117-138, 1997

LARVAE OF FIVE FISH FAMILIES WITH PATfERN 10 OF THE RAMUS LATERALIS ACCESSORIUS NERVE (ARRIPIDAE, GIRELLIDAE, KYPHOSIDAE, MICROCANTHIDAE AND SCORPIDIDAE): RELEVANCE TO RELATIONSHIPS

Francisco J. Neira, Anthony G. Miskiewicz and Barry D. Bruce

ABSTRACT We describe the larvae of five temperate Australian fishes from five percoid families considered to form part of a monophyletic group because they share the uncommon pattern 10 of the facial nerve ramus lateralis accessorius (RLA-IO), An attempt was made to assess to what extent the larval characters described for these species-Arripis trulta (Arripidae), Gi- rella tricuspidata (Girellidae), Kyphosus sp. (Kyphosidae), Atypichthys strigatus (Microcan- thidae) and Scorpis (lineolata?) (Scorpididae)-support a close relationship among these RLA-lO families, Larvae of the five species share similarities in body form, head spination, fin formation and pigment pattern, and generally fit the "distinctive larval form" previously described for "girelloid" RLA-IO families. Although such similarities are consistent with a close relationship among these families, most attributes of the distinctive larval form are also present in larvae of a number of non RLA-1O percoid families, notably the Pomatomidae. Based only on larval characters, one could hypothesize that most RLA-1O percoid families, including the Dichistiidae (Leis and van der Lingen, 1997), form part of a larger group that also includes non RLA-IO families such as the Pomatomidae. Alternatively, the "distinctive larval form" in Pomatomus and some other non RLA-lO families could have been acquired independently. However, since we could not show that any of the attributes of the distinctive larval form constitute synapomorphies, nor is the sister group of the RLA-IO families known, neither of these alternatives could be tested during this study. Larval characters alone cannot at present resolve the relationships of the RLA-1O families.

The Percoidei is a highly diverse and probably polyphyletic suborder of perciform fishes (Johnson, 1984, 1993; Johnson and Patterson, 1993; Nelson, 1994). Eight percoid families, namely the Arripididae, Girellidae, Kyphosidae, Khuliidae, Oplegnathidae, Microcanthidae, Scorpididae and Terapontidae, and the six families of the suborder Stromateoidei, are considered to form a monophyletic group based on their possession of the uncommon pattern 10 of the facial nerve ramus lateralis accessorius (RLA-lO). This character, distinct from the nerve pattern 9 shared by most percoid fishes, is considered a synapomorphy uniting most groups (excluding the carangoid Nematistius which possesses a reduced RLA-lO pattern) that possess it (Freihofer, 1963; Johnson and Fritzsche, 1989). The Girellidae, Kyphosidae and Scorpididae (GKS herein) are generally considered to be closely related but separate families (Johnson, 1984, 1993; Nelson, 1994), although they have been variously classified as subfamilies of the Ky- phosidae or Girellidae (Greenwood et aI., 1966). Their recognition as closely related families was indicated by Johnson (1984) and latter supported by Johnson and Fritzsche (1989) and Stevens et aI. (1989) based on the overall similarity of their larvae. The position of the Microcanthidae is less clear. Microcanthids have been included in the Chaetodontidae (Fraser-Brunner, 1945), Scorpididae (Springer, 1982; Last et aI., 1983; Kojima, 1988; Gomon et aI., 1994) and Kyphosidae (Nelson, 1994). Although microcanthids are considered to be closely related to the GKS families by the common possession of the RLA-I 0 (Johnson and Fritzsche, 1989),

117 118 BULLETIN OF MARINE SCIENCE, VOL. 60, NO.1, 1997

Johnson (1984) had previously stated that, based on larval characters, evidence to support this relationship was lacking. However, his observation was based on a larval series illustrated in Uchida et aI. (1958) as Microcantus strigatus but which in fact contained a larva (Uchida et aI., 1958: pI. 60, fig. 4) of the terapontid Terapon jarbua (Leis and van der Lingen, 1997). Larvae of some representatives of the Girellidae (Girella), Kyphosidae (Her- mosilla, Kyphosus), Microc:anthidae (Microcanthus) and Scorpididae (Labraco- glossa, Medialuna) are known (Table 1). The purpose of this paper is to provide descriptions and illustrations of larvae of additional RLA-lO taxa not previously described and to assess whether they possess the "distinctive larval form" that led Johnson (1984) and others to propose a close relationship among the GKS (= "girelloid") families. We include the first description of a member of the family Arripidae (Arripis trutta), and the first of a member of the microcanthid genus Atypichthys and the scorpidid genus Scorpis. We also describe the larvae of one species of each Girella and Kyphosus. Descriptions of the five taxa treated in this paper will appear elsewhere (Neira et aI., 1997). We limited our comparisons to the four main character groups which were regarded by Stevens et al. (1989) and Johnson and Fritzsche (1989) as similar in GKS larvae: body morphology, head spination, fin development and pigment pattern. We also extended our comparisons to larvae of the other three percoid RLA-lO families Kuhliidae, Oplegnath- idae and Terapontidae which have been well described (Leis and Rennis, 1983; Johnson, 1984; Kinoshita, 1988a, 1988b, 1988c; Leis and Trnski, 1989).

MATERIALS AND METHODS

Material Examined.-Larvae were obtained from samples collected in estuarine and coastal waters of temperate Australia, mainly off the central New South Wales coast and northeastern Tasmania. Larval collections were supplemented with dipnetted and other museum specimens when available, The total number of specimens examined, size range (mm) and collection locality are provided for each taxa. Representatives of all taxa examined are lodged in the Australian Museum, Sydney. Terminology of developmental stages, head spines and morphological measurements follow Leis and Trnski (1989). All measurements were made to the nearest 0.1 mm using an ocular micrometer fitted to a stereomicroscope. Body length (BL) corresponds to notochord length (tip of snout to tip of notochord) in preflexion and flexion larvae and to standard length (tip of snout to posterior margin of hypurals) in postflexion larvae. Measurements of body depth (BD), head length (HL) and preanal length (PAL) given throughout the text and in Table 2 are expressed as a percentage of body length. We followed Leis and Trnski (1989) in using body depth and head length to describe a larva as either elongate (BD = 10-20% BL) or moderate (BD = 20-40% BL) and as having a head either moderate (HL = 20-33% BL) or large (HL >33% BL). In addition, we used preanal length to characterise a larva as having a gut either moderate (PAL = 30-50% BL) or long (PAL = 50-70% BL). Pigment refers solely to melanin, All larvae were illustrated with the aid of a drawing tube attached to the stereomicroscope,

Identification and Format of Descriptions.-Larvae were identified to the lowest possible taxon using literature descriptions (see references in Table 1) and the size series method (Leis and Trnski, 1989). A developmental series was assembled for each species using body morphology, head spines, gut length and shape, sequence of fin development, and pigment patterns. The largest specimen in each of the series was identified to genus and/or species using meristic counts obtained from the literature. Information on adult distributions and locations of larval collections facilitated the identifications. Since the main objective of our paper is to evaluate possible relationships among the five RLA-1O families using external larval characters of representative taxa, our individual descriptions do not follow the dynamic approach that we would normally take when describing larval development in detail. Instead, we provide descriptions in a telegraphic style and summarise in tables many of the developmental characters normally used in full descriptions, induding morphometries, notochord flexion, number of myomeres and sequence of fin formation (Table 2). Because descriptions of the five taxa treated here are also given in the forthcoming publication by Neira et al. (1997), relevant references are provided in each case. NEIRA ET AL.: LARVAE OF FIVE AUSTRALIAN FISHES 119

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RESULTS

Descriptions of Larvae Arripis trutta (Bloch and Schneider, 1801)-Arripidae Figure 1

18 specimens, 2.7-19.7 mm-coastal waters off Sydney and northeastern Tasmania Late postflexion larvae were identified as Arripis by the dorsal (IX, 13-19) and anal fin counts (III, 9-10) and as A. trutta by the number of gill rakers along the first gill arch (9-13 + 20-24) (Paulin, 1993; Gomon et aI., 1994). Larvae are also described in Bruce et al. (1997a). Morphology.-Body moderate; head round, moderate to large (Table 2). Two small spines on outer preopercular margin from flexion, up to seven by postflexion and in early juveniles (Fig. IB, D, E); spines gradually reduce in size, forming a serrate preopercle by 19.7 mm. One to three small spines on inner preopercular margin and one supracleithral spine by early postflexion (Fig. ID). Low, smooth supraocular ridge from early flexion. One opercular, posttemporal, interopercular and cleithral spine and two supracleithral spines in early juveniles (Fig. IE). Gas bladder visible over foregut when inflated. Gut initially straight, coiled and compact by 3.2 mm, moderate to long (Table 2). Prominent gap between anus and origin of anal fin in early stages closes by late postflexion. Pigmentation.-Early larvae are moderately pigmented, heavily pigmented from early postflexion (Fig. I). Head pigment in preflexion and flexion larvae includes several melanophores externally over midbrain, one or two internally below otic capsule, small melanophores at tip of upper jaw, a large internal melanophore under nasal pit and a conspicuous melanophore on inner surface of upper preopercle. Additional melanophores dorsally over head, along dentary, ventrally anterior to cleithral symphysis, and over and under opercle following flexion. Heavy pigment on head in early juveniles (Fig. IE). Moderate internal pigment over peritoneum, dorsally over gas bladder and gut, and external melanophores laterally over gut in preflexion and flexion larvae (Fig. lA, B); heavy pigment over entire gut in early juveniles (Fig. IE). Distinct pigment dorsally on trunk and over dorsal, lateral and ventral surfaces of tail in larvae prior to juvenile stage (Fig. lA-D). Four to six internal melanophores on trunk, anterior-most on nape, followed by a series of external melanophores along dorsal midline of tail in preflexion and flexion larvae, from myomeres 15-16 to notochord tip (Fig. IA-C); internal trunk melanophores obscured by heavy dorsal pigment in early juveniles (Fig. ]E). Prominent melanophore series along dorsolateral surface of trunk by early flexion, extending posteriorly onto tail and almost reaching end of dorsal-fin base by early postflexion. Three to six external melanophores along lateral midline of tail by late preflexion, between myomeres 16 and 22, and an internal series ventrally along notochord visible below external series (Fig. IA); both series extend anteriorly, the external series forming a prominent lateral stripe by early postflexion (Fig. ID). Few, widely spaced internal melanophores and a series externally along ventral midline of tail to end of caudal peduncle; melanophores along anal-fin base internal by early postflexion. Pigment on membrane between dorsal-fin spines and heavy pigment over entire trunk and tail, except for pectoral-fin base and part of caudal peduncle, in early juveniles (Fig. IE). Prominent melanophores around notochord tip, and pigment under posterior 122 BULLETIN OF MARINE SCIENCE, VOL. 60, NO. I, 1997

A . "

Figure 1. Larvae and early juvenile of the arripid Arripis trutta from northeastern Tasmania, (A) 3.7 mm BL preflexion; note pigment surrounding notochord tip. (B) 4.9 mm BL flexion, with developing pelvic-fin bud. (C) Dorsal view of larva in B showing series of internal melanophores along nape. (D) 5.7 mm BL early postflexion; dorsal and anal fins still developing. (E) 8,3 mm BL early juvenile. Illustrated by F. J. Neira (adapted from Bruce et aI., 1997a). NEIRA ET AL.: LARVAE OF FIVE AUSTRALIAN FISHES 123

notochord extending ventrally to hypural region, in preflexion and flexion larvae (Fig. 1). Few melanophores remain around notochord tip by postflexion; pigment below posterior notochord persists between upper and lower hypural plates during postflexion, with additional melanophores along caudal-fin base.

Girella tricuspidata (Quoy and Gaimard, 1842)-Girellidae Figure 2

25 specimens, 3.0-12.4 mm-NSW coastal waters

Late postflexion larvae were identified as G. tricuspidata by the dorsal (XIV- XVI, 11-13) and anal (III, 11-12) fin counts (Kuiter, 1993; Gomon et al., 1994). Dorsal fin-ray counts can be used to distinguish this species from co-occurring G. elevata (XIII, 14). Larvae G. tricuspidata are also described in Miskiewicz (1987) and Miskiewicz and Trnski (1977).

Morphology.-Body moderate; head round and moderate (Table 2). One small spine on inner preopercular margin from preflexion and up to two and three small spines on inner and outer preopercular margins by postflexion, respectively (Fig. 2A, C, D). Smooth supraocular ridge by early postflexion and one supracleithral and opercular spine in postflexion larvae >7.8 mm. Gas bladder not visible. Gut coiled and compact, moderate to slightly long (Table 2). Prominent gap between anus and origin of anal fin persists to late postflexion (Fig. 2D) and closes by settlement.

Pigmentation.-Larvae are lightly to moderately pigmented. Head pigment in preflexion larvae consists of one large and two small external melanophores above midbrain and a band of internal pigment from under forebrain to below otic region (Fig. 2A). One large melanophore over forebrain, one on opercle and one internal melanophore laterally on hindbrain by flexion (Fig. 2B); additional melanophores on jaw tips, ventrally anterior to c1eithral symphysis, and one at angle of lower jaw by postflexion. Moderate pigment on head by postflexion (Fig. 2D). Internal pigment over peritoneum and dorsally over gas bladder and gut in all stages. Two external melanophores ventrally and several laterally on gut in preflexion and flexion larvae (Fig. 2A, B). Heavy pigment over entire gut by postflexion (Fig. 2D). Characteristic pigment dorsally on trunk and on dorsal, lateral and ventral midlines of tail in all stages (Fig. 2). One large internal melanophore on nape and three to six large, widely spaced melanophores along dorsal midline of tail in preflexion and flexion larvae (Fig. 2A, B). External melanophore series along posterior lateral midline of tail from late preflexion, with an internal series dorsally and ventrally along posterior notochord (Fig. 2B); additional external melanophores anteriorly along lateral midline forming a prominent lateral stripe by postflexion (Fig. 2D). Five to eight large, widely spaced melanophores along ventral midline of tail in preflexion and flexion larvae. Additional external melanophores dorsally along trunk and tail and ventrally along tail by 10.0 mm. One or two internal melanophores under notochord tip and one elongate melanophore in hypural region in preflexion larvae (Fig. 2A). Melanophores under tip disappear during flexion; elongate melanophore remains at base of lower caudal-fin rays from postflexion (Fig. 2C). Pigment along upper hypural margin in postflexion larvae. 124 BULLETIN OF MARINE SCIENCE, VOL. 60, NO. I, 1997

Figure 2. Larvae of the girellid Gire/La tricuspidata from central New South Wales. (A) 4.1 mm BL preflexion. (B) 5.0 mm BL flexion; note developing dorsal and anal fins, (C) 6,8 mm BI early postflexion; note low, smooth supraocular ridge. (D) 12.4 mm BL poslflexion; note small supracleithral spine. Illustrated by T. Trnski (adapted from Miskiewicz and Trnski, 1977).

Kyphosus sp.-Kyphosidae Figure 3

28 specimens, 2.1-10.8 mm--NSW coastal waters Late postflexion larvae were identified as Kyphosus sp. following Walker and Watson (1983) and by the dorsal (XI, 12-16) and anal (III, 10-11) fin counts NEIRA ET AL.: LARVAE OF I'IVE AUSTRALIAN FISHES 125

Figure 3. Larvae of the kyphosid Kyphosus sp. from central New South Wales. (A) 2.7 mm BL preflexion. (B) 4.1 mm BL late preflexion, with developing dorsal and anal fins. (C) Dorsal view of larva in B showing series of internal melanophores along nape. (D) 5.2 mm BL late flexion; note developing pelvic-fin bud. (E) 6.1 mm BL postflexion. Illustrated by E J. Neira (adapted from Neira and Miskiewicz, 1997). 126 BULLETIN OF MARINE SCIENCE, VOL. 60, NO. I, 1997

(Kuiter, 1993; Gomon et aI., 1994). Larvae are also described in Neira and Mis- kiewicz (1997). Morphology.-Body moderate; head round, moderate to large (Table 2). Two or three small spines on outer preopercular margin and a smooth supraocular ridge and a posttemporal ridge by flexion (Fig. 3D). One spine on inner and up to seven along outer preopercular margin, one supracleithral and a weak opercular spine by early postflexion (Fig. 3E); preopercular spines reduce in size by late postflexion. Small gas bladder above foregut. Gut straight in yolk-sac larvae and loosely coiled from late preflexion, long in all stages (Table 2). Preanal membrane through to postflexion. No gap between anus and origin of anal fin. Pigmentation.-Early larvae are moderately pigmented, heavily pigmented from postflexion (Fig. 3). Head pigment in prefiexion and flexion larvae comprises several melanophores dorsally over midbrain, none to few at tip of jaws and on snout, one melanophore anterior to cleithral symphysis, one distinct on upper preopercle, one internally at roof of mouth, and pigment in otic region and under hindbrain (Fig. 3A, B). Additional melanophores dorsally over head, several scattered over snout, along jaws and opercular area by postflexion (Fig. 3E). Internal pigment over peritoneum and dorsally over gas bladder in all stages (Fig. 3). Several internal melanophores dorsally along hindgut, and three to five external melanophores ventrally on gut in preflexion and flexion larvae. Addi- tional melanophores laterally on gut during flexion and heavy pigment over entire gut by postflexion (Fig. 3E). Characteristic pigment dorsally on trunk and over dorsal and ventral surfaces of tail in all stages (Fig. 3). Four semi-internal (=just under surface) melanophores on trunk in preflexion larvae, anterior-most on nape, followed by 11-12 large, stellate melanophores along dorsal midline between myomeres 7 and 22 (Fig. 3A); nape and next three trunk melanophores become entirely internal during flexion (Fig. 3B, C). Scattered melanophores laterally over trunk and tail, except caudal peduncle, from late flexion; distinct dark stripe posteriorly along lateral midline of tail by postflexion (Fig. 3E). Series of 12-13 melanophores along ventral midline of tail in preflexion larvae, gradually becoming internal during flexion. Heavy pigment over entire body in postflexion larvae >8 mm, including caudal peduncle and all fins. Several small melanophores under notochord tip in preflexion larvae (Fig. 3A); these become internal during flexion and most disappear by postflexion except for one external and one internal melanophore that persist at junction of hypural plates (Fig. 3E). Pigment along caudal-fin base from flexion.

Atypichthys strigatus (Gunther, 1860)-Microcanthidae Figure 4

24 specimens, 2.9-14.4 mm--NSW coastal waters Late postflexion larvae were identified as A. strigatus by the dorsal (XI-XII, 16-18) and anal (III, 15-17) fin counts (Gomon et al., 1994), and distinguished from that of the co-occurring Microcanthus strigatus using pigmentation patterns (Uchida et aI., 1958; Walker, 1983; Kojima, 1988). Larvae A. strigatus are also described in Miskiewicz and Neira (1997). Morphology.-Body slightly elongate to moderate; head round and moderate (Table 2). Two small spines on inner and two on outer preopercular margin from late preflexion-early flexion; spines increase in number on both margins, those on outer margin becoming longer by late flexion but reducing in length thereafter (Fig. 4B- NEIRA ET AL.: LARVAE OF FIVE AUSTRALIAN FISHES 127

Figure 4. Larvae of the microcanthid Atypichthys strigatus from central New South Wales. (A) 2.9 mm BL preflexion; note pigment surrounding notochord tip. (B) 6.4 mm BL late flexion; note developing pelvic-fin bud. (C) 9.8 mm BL postflexion. (D) 14.4 mm BL late postflexion. Illustrated by F. J. Neira (adapted from Miskiewicz and Neira, 1997). 128 BULLETIN OF MARINE SCIENCE. VOL. 60, NO. I. 1997

D). Smooth supraocular ridge from early flexion, serrate from late flexion. One subopercular and a supracleithral spine by late flexion (Fig. 4B); supracleithral spine strongly serrate by postflexion (Fig. 4D). Additional interopercular,cleithral and posttemporal spines by postflexion. Small gas bladder above foregut. Gut coiled and compact, moderate to slightly long (Table2). Prominent gap between anus and origin of anal fin in early stages closes by late postflexion (Fig. 4D). Pigmentation.-Early larvae are lightly pigmented, moderately pigmented by postflexion (Fig. 4). Head pigment in preflexion larvae consists of one to three large melanophores over midbrain and pigment under otic capsule. Several melanophores internally in nostrils and under opercle, and several externally over forebrain, jaw tips, ventrally anterior to cleithral symphysis, and a prominent melanophore just below supracleithral spine during flexion (Fig. 4B, D). Heavy pigment on head in postflexion larvae, before pigment develops laterally on trunk and tail (Fig. 4C, D). Internal pigment dorsally over gas bladder and gut in all larvae, becoming heavier by postflexion. Pigment over peritoneum and one or two small melanophores ventrally on gut in preflexion larvae, one ventrally at anus; latter disappears by postflexion. Numerous external melanophores laterally over gut during flexion; pigment over entire gut by postflexion. One melanophore on outer margin of pectoral-fin base during flexion, several by postflexion. Distinct pigment dorsally on trunk and on dorsal and ventral surfaces of tail in all stages (Fig. 4). Five large, widely spaced melanophores along dorsal midline of trunk and tail in preflexion larvae; anterior-most at nape (Fig. 4A); latter be- comes internal by late pn~f1exionwhile remaining four persist through to postflexion spaced along dorsal-fin base. Internal melanophore series above posterior notochord in flexion larvae, followed by an external series along lateral midline of trunk and tail in postfIexion larvae ca. 10 mm. Three to four large, widely spaced melanophores along ventral midline of tail in preflexion larvae, remaining spaced along anal-fin base from late flexion (Fig. 4). Melanophores over spinous portion of dorsal fin and over pelvic fins by late postflexion (Fig. 4D). One melanophore dorsally and ventrally over caudal peduncle and pigment surrounding notochord tip in preflexion larvae (Fig. 4A); both melanophores on caudal peduncle become internal before flexion and remain around urostyle in postflexion larvae. Dorsal pigment disappears during flexion while pigment under notochord tip remains along caudal-fin base by postftexion.

Scorpis (lineolata?) Kner, 1865-Scorpididae Figure 5

28 specimens. 4.3-13.3 mm--NSW coastal waters Late postftexion larvae were identified as belonging to the Scorpididae following Konishi (1988a) and tentatively as Scorpis (lineolata?) by the dorsal (IX, 27) and anal (TIl, 28) fin counts, and the fact that it is the only common Scorpis species in temperate eastern Australia (Kuiter, 1993; Gomon et aI., 1994). Larvae are also described in Rissik et aI. (1997). Morphology.-Body moderate; head round, moderate to slightly large (Table 2). Two spines on inner preopercular margin during flexion (Fig. 5B), only one remaining by postflexion, small and blunt. Two small spines on outer preopercular margin in late preflexion larvae, up to eight in late postflexion larvae (Fig. 5D). Smooth supraocular ridge and one interopecular spine from late flexion. Two small subopercular, three interopercular, two posttemporal and a cleithral and su- NEIRA ET AL.: LARVAE OF FIVE AUSTRALIAN FISHES 129

Figure 5. Larvae of the scorpidid Scorpis (lineo/ata ?) from central New South Wales. (A) 5.0 mm BL preflexion; note pigment surrounding notochord tip. (B) 6.2 mm BL flexion, with developing dorsal and anal fins. (C) 7.7 mm BL late flexion; note developing pelvic-fin bud. (D) 8.6 mm BL postflexion. Illustrated by F. J. Neira (adapted from Rissik et aI., 1997).

pracleithral spine in postflexion larvae (Fig. 5D). Small gas bladder above foregut, obscured by postflexion due to heavy pigment. Gut coiled and compact, moderate to slightly long (Table 2). Prominent gap between anus and origin of anal fin closes by postflexion (Fig. 5D). 130 BULLEHN OF MARlNE SCIENCE, VOL. 60, NO. ], ]997

Pigmentation.-Early larvae are moderately pigmented, heavily pigmented by postflexion. Head pigment in preflexion and flexion larvae consists of several external melanophores dorsally over brain and jaw tips, and internal pigment under brain and over hindbrain (Fig. 5A, B). Two to three melanophores anterior to cleithral symphysis from flexion and a distinct patch of large, stellate melanophores under eye from late flexion. Heavy pigment on head from late flexion, before pigment develops laterally on trunk and tail (Fig. 5C). Heavy internal pigment dorsally over gas bladder and gut in all stages. Three to four external melanophores ventrally along gut in preflexion and flexion larvae. Several melanophores laterally over gut from flexion and heavy pigment over entire gut by early postflexion (Fig. 5C, D). Distinct pigment dorsally on trunk and on dorsal and ventral surfaces of tail in all stages (Fig. 5). Two internal melanophores on nape followed by 8-13 large, closely-spaced melanophores along dorsal midline of trunk and tail in preflexion and flexion larvae (Fig. 5A, B). External melanophore series along posterior midline of tail in preflexion and flexion larvae, with an internal series along notochord visible above and below external series; extemal series extends anteriorly forming a continuous stripe along lateral midline by postflexion (Fig. 5C, D). Additional series of internal melanophores above anterior notochord from flexion, extending posteriorly with growth (Fig. 5B, C). Up to 13 large melanophores extending upwards internally along anterior ventral midline of tail in preflexion and flexion larvae, followed by a continuous series of small melanophores along posterior tail region to caudal peduncle; anterior melanophores become entirely internal by late flexion, while posterior melanophores become discrete by early postflexion. Pig- ment from dorsal and lateral midlines extends over trunk and tail and towards head from late flexion. Heavy pigment dorsally and laterally over body by postflexion, except for a small region of caudal peduncle; little or no pigment along ventral surface of trunk and tail apart from a few melanophores at distal end of anal fin (Fig. 5D). Several melanophores surrounding notochord tip and one elongate melanophore on hypural region in preflexion and flexion larvae (Fig. 5A, B). Melanophores above notochord tip disappear during flexion; melanophores under notochord tip and elongate hypural melanophore remain along caudal-fin base in postflexion larvae (Fig. 5D).

DISCUSSION

Family Relationships Based on Adult Characters.-All five families treated in this study possess the uncommon RLA-lO pattern (Freihofer, 1963), which was interpreted by Johnson and Fritzsche (1989) as a synapomorphy uniting most percoid families possessing it (Freihofer [1963] did not consider scorpidids to have RLA-lO pattern but G. D. Johnson and J. M. Leis [pel's. comm.] found a somewhat modified pattern 10 in material of this family they examined). Johnson and Fritzsche (1989) found no other convincing synapomorphies uniting the families Girellidae (G), Kyphosidae (K) and Scorpididae (S). In fact, apart from their general external similarity, the small, nibbling-type mouth (Gosline, 1985) and possibly the progenic serial replacement of primary teeth, Johnson and Fritzsche (1989) concluded that all osteological characters shared by these families were either widespread percoid symplesiomorphies or were not unique to the GKS families. Relationships between the Arripidae and Microcanthidae, both RLA-lO families, and between these and the GKS families have not been assessed and a NEIRA ET AL.: LARVAE OF FIVE AUSTRALIAN FISHES 131 comparative examination of adult characters to establish relationships among these families is beyond the scope of this paper. Family Relationships Based on Larval Characters.-A close relationship between the "girelloid" families GKS was tentatively proposed by Johnson and Fritzsche (1989) from the evidence on larval characters presented by Stevens et al. (1989). The latter authors observed that larval Girella nigricans (Girellidae), Hermosilla azurea (Kyphosidae) and Medialuna californiensis (Scorpididae) from California shared similarities in body morphology, head spination and pigment patterns. Fol- lowing Stevens et al. (1989), Johnson and Fritzsche (1989) regarded the GKS families as having a "distinctive larval form" characterised by strong similarities in four main aspects: (a) body form; (b) generalised head spination; (c) pattern of fin development; and (d) pigment pattern. It is worth noting, however, that Johnson (1984) had earlier mentioned a possible close relationship between gi- rellids and scorpidids based on their similar larvae. Leis and van der Lingen (1997) added the RLA-lO families Microcanthidae (M) and Dichistiidae (D) to the group possessing the distinctive GKS larval form (=GKS + MD). In many aspects, our GKMS larvae and those of the previously undescribed Arripis trutta (Arripidae = A) fit the pattern of the distinctive GKS + MD larval form. Indeed, all our AGKMS larvae described in this study possess most attributes of the distinctive GKS larval form of Johnson and Fritzsche (1989), including the head spination and pigment patterns. However, as clearly pointed out by Johnson and Fritzsche (1989), Stevens et al. (1989) and Leis and van der Lingen (1997), not one of the characters that constitute the distinctive larval form is by itself exclusive to the above RLA-lO families and certainly none is unique among percoid larvae (Leis and Rennis, 1983; Leis and Trnski, 1989). To assess to what extent larval characters reflect relationships among the RLA-lO families considered here, we firstly describe each of the characters that comprise the distinctive GKS larval form and then compare them with those of our AGKMS larvae. BODYFORM.The distinctive GKS larval form, as described by Johnson and Fritzsche (1989) and Leis and van der Lingen (1997), is characterised by a laterally compressed body, a rounded head, a coiled, compact gut and a prominent gap between anus and origin of anal fin. Stevens et al. (1989) noted that a major morphological difference among the larvae of the GKS species was an increase in the degree of body robustness from Girella to Medialuna to Hermosilla. A similar pattern to that of the Californian species was observed in our AGKMS larvae when considering the late postflexion stage, with Girella being the least robust and then increasing in robustness through to S, A, M and K. A rounded head was also present in all our AGKMS larvae. The gut length and condition, and the persistence of the gap between anus and origin of anal fin vary amongst our AGKMS larvae. Larvae of AGMS have a moderate gut in the early stages (PAL 30-50% BL), becoming moderately long (PAL >50% BL) following flexion, whereas it is always long in Kyphosus larvae (PAL 51-69%). Similarly, the gut is tightly coiled and compact in all except Kyphosus sp. larvae, where it is loosely coiled (Table 2). The gap between anus and origin of anal fin is initially large in early AMS larvae but closes after anal- fin formation is complete. By contrast, a prominent gap is retained in Girella larvae until just prior to settlement whereas there is no gap in Kyphosus sp. larvae. GENERALISEDHEADSPINATlON.Johnson (1984) considered the distinct GKS larval form as having a "generalised head spination," comprised primarily of a series of weak spines on both margins of the preopercle, and other weak spines 132 BULLETJN OF MARINE SCIENCE, VOL. 60, NO. I, 1997 such as subopercular, interopercular, posttemporal and supracleithral in varying numbers. Preopercular and supracleithral spines are present in all our AGKMS larvae and range from weak to moderate. Head spination was least developed in Girella, which had only a few small preopercular spines and a supracleithral spine, and was most elaborate in Atypichthys, which had the most extensive preopercular spination, including a moderately elongate spine at the angle in flexion and early postflexion larvae, and also a serrate supraocular ridge, and a prominent, serrate supracleithral ridge (Table 2).

SEQUENCEOF FIN DEVELOPMENT.The most common sequence of fin development in percoid fishes starts with the caudal fin (C), followed by the dorsal and anal fins simultaneously (D, A), then the rays of the pectoral fin (PJ) and finally the pelvic fin buds (P2) (Johnson, 1984). This sequence of fin development was observed in all our GKMS except Arripis larvae, where the P2 buds begin to develop prior to the PI rays (Table 2). PIGMENTATION.The overall pigmentation pattem has been regarded as one of the distinguishing features of the distinctive GKS larval form (Johnson and Fritz- sche, 1989; Stevens et al. 1989). Leis and van der Lingen (1997) identified nine pigment character groups within GKMS larvae. These were (1) dorsal midline series on tail; (2) ventral midline series on tail; (3) lateral midline series on tail; (4) internal series along vertebrae; (5) pigment on caudal anlage/hypural edge; (6) anterior progression of pigment on trunk and tail; (7) internal pigment stripe from snout to opercle through the eye; (8) early-forming, heavy pigment on brain and nape; and (9) heavy pigment dorsally and laterally on gut. All but character (6) were present in various degrees in our AGKMS larvae (Table 3). Another pigment character found to be relevant to the distinctive GKS larval form in this study was the presence of pigment on the notochord tip area in all preflexion and flexion AGKMS larvae, which is probably included in character (5) of Leis and van der Lingen (1997). Many of these pigment characters also occur in various combinations in larvae of non RLA-lO families and therefore are of limited use for assessing relationships but very useful in distinguishing between species. Before flexion, overall pigmentation ranges from light in our GM larvae to moderate in AKS larvae. Soon after flexion, pigment is heavy over the entire body in AKS larvae, increasing in intensity from Kyphosus to Scorpis to Arripis. Pigment in postflexion larvae of Girella and Atypichthys remains light and mostly restricted to the dorsal, lateral and ventral midlines of the trunk and tail, and does not develop over the rest of the trunk and tail until the beginning of the juvenile stage. In addition, the timing of development of pigment in some areas varies considerably among speci4~s(Table 3). For example, head pigment forms either simultaneously with (A larvae), mostly after (K larvae) or before (GMS larvae) that on the trunk and tail Cfable 3). Pigment along the dorsal and ventral midlines of the trunk and tail (characters 1 and 2) is present in all our AGKMS larvae and occurs in two general patterns (Table 3): a few large, discrete melanophores spaced along both midlines as in the GM larvae, or a continuous melanophore series along the dorsal midline, with a variable pattern of internal and external melanophores along the ventral midline, as in the AKS larvae. In addition to this dorsal pigment, all our AGKMS larvae have pigment on the nape (character 8 in part): Girella and Atypich.thys each have one internal melanophore on the nape, initially external in the latter, Scorpis has two, and Arripis and Kyphosus have one, closely followed by three to five internal trunk melanophores (Figs. IC, 3C; Table 3). NEIRA ET AL.: LARVAE OF FIVE AUSTRALIAN FISHES 133

, to Ol E "'0 II.l () c..to '"• '"II.l II.l •••• bIlO _c..a..c """I 0!':: C')

o!':: ";;: II.l a:: •...II.l c.. II.l IS 134 BULLETIN OF MARINE SCIENCE. VOL 60. NO. I. 1997

Pigment along the lateral midline forms early during development in AGS larvae and consists of a series of external melanophores on the posterior region of the tail (character 3), and another series of internal melanophores over and/or under the notochord (character 4) that forms roughly at the same time and in the same area as the external series. By contrast, internal pigment along the notochord is absent in Kyphosus larvae, is present but late to form in Atypichthys larvae, and both taxa develop an external midlateral tail series only in the postflexion stage. Pigment is always present on the notochord tip area (character 5) in preflexion and early flexion AGKMS larvae, and occurs in two patterns: either around or under the tip. Pigment around the tip is present in AMS larvae and increases in intensity from Atypichthys, to Scorpis and to Arripis. Pigment under the tip occurs in GK larvae, comprising a single melanophore in Girella and several in Kypho- sus. The pattern in our GKS larvae was almost identical to that of the Californian GKS larvae described by Stevens et al. (1989), with one melanophore under the tip in G. nigricans (newly-hatched larvae also have dorsal melanophores but these disappear prior to flexion), and several melanophores under the tip in H. azurea and around the tip in M. californiensis.

Comparisons with Other RIA-JO Families.-Larvae of other RLA-lO families possess most attributes of the distinctive GSK larval form although in different combinations and varying degrees of development. For example, kuhliid, oplegnathid and terapontid larvae all have a rounded head, tightly coiled, pigmented guts, and a temporary gap between the anus and the origin of the anal fin which is prominent in kuhliids and terapontids and small in oplegnathids (Leis and Rennis, 1983; Kinoshita, 1988a, 1988b, 1988c; Leis and Trnski, 1989). Sequence of fin development in larvae of these families also follows the common percoid pattern, i.e., C-D,A-PI-P2• Head spination is also generalized in the three families, and weak to moderate except perhaps in some terapontids (Johnson, 1984; Ki- noshita, 1988a, 1988c; Leis and Trnski, 1989; Trnski and Neira, 1997). Most of the pigment characters of the distinctive larval form are also present in larvae of these families, although the pattern may differ markedly. Trunk and tail pigment in early kuhliid, oplegnathid and terapontid larvae is restricted to the ventral midline and is generally not as prominent as in our AGKMS larvae. In addition, pigment along the dorsal midline in larvae of these families is lighter and usually develops during the postflexion stage (Kinoshita, 1988a, 1988b, 1988c; Leis and Trnski, 1989). Some terapontid larvae (e.g. Terapon jarbua) possess an external/internal melanophore pattern along the posterior lateral midline similar to that in our early AGS larvae but this forms by late postflexion stage (Kinoshita, 1988b; Leis and Trnski, 1989). Such early-forming pattern is absent in kuhliid and oplegnathid larvae although both have internal melanophores dorsally along the vertebrae in the postflexion stage, and only kuhliid larvae develop a distinct external lateral stripe (Kinoshita, 1988a, 1988c; Leis and Trnski, 1989). All except oplegnathid larvae have pigment associated with the notochord tip: few melanophores restricted to under the tip in terapontids, and a few melanophores over the tip and a distinctive prominent melanophore cluster in the hypural region in kuhliids (Kinoshita 1988a, 1988b; Leis and Trnski, 1989; Trnski and Neira, 1997). The RLA-lO pattern also occurs in the Stromateoidei (Freihofer, 1963; Johnson and Fritzsche, 1989). Larvae of the stromateoid family Centrolophidae, considered to be near the base of the suborder (Hom, 1984), also possess many of the attributes of the distinctive AGKMS larval form, including 25-26 myomeres, a round- NEIRA ET AL.: LARVAE OF FIVE AUSTRALIAN FISHES 135 ed head, weak head spination comprising preopercular and subopercular spines, a coiled and pigmented gut, and pigment along the dorsal, lateral and ventral midlines of tail (Bruce et aI., 1997b). Relationships within the Stromateoidei are considered by Haedrich (1967) and Hom (1984), and hence, are not dealt with in this study. Comparison with non RLA-10 Percoid Families.-Larvae of many non RLA-lO percoid families share some to many of the characters of the distinctive larval form with the AGKMS families. Examples include some carangids, cheilodactylids, emmelichthyids, gerreids, leptobramids, mullids, and some haemulids, po- macentrids and pomatomids (Leis and Rennis, 1983; Johnson, 1984; akiyama, 1988; Leis and Trnski, 1989). Of these, larvae of the pomatomid Pomatomus saltatrix possess all features that characterise the distinctive AGKMS larval form: 25-26 myomeres; a rounded head; a coiled, compact and pigmented gut; a small gap between anus and origin of anal fin which closes during flexion; weak head spination including preopercular and opercular spines, and weak supraocular and posttemporal ridges; and pigment over the head and nape, and along the dorsal, lateral and ventral midlines of tail (Trnski et al., 1977). Cheilodactylid and lep- tobramid larvae also possess some of the characters of the distinctive larval form including a rounded head, a pigmented, coiled gut and similar pigment patterns (Bruce, 1989; Leis and Trnski, 1989). Specific pigment characters which occur in some of our AGKMS larvae are also present in larvae of some other non RLA-lO families. For example, the early-forming, external/internal pigment pattern along the posterior lateral midline in Arripis, Girella and Scorpis larvae also occurs in some pomacentrid larvae, particularly of Chromis and Pomacentrus (Leis and Rennis, 1983; Kinoshita, 1988d). Larvae of the AGKMS species described in this study are neustonic or occur in surface waters (Miskiewicz, 1987; Gray 1993; Bruce et al., 1997a) and thus possess the heavy pigment (in later stages, at least) that characterises the larvae of many species living in these habitats, including a variety of non RLA-I 0 families (e.g., cheilodactylids, carangids and pomatomids; Leis and Rennis, 1983; Bruce, 1989, Leis and Trnski, 1989, Trnski et aI., 1977). As suggested by Leis and van der Lingen (1997), however, this heavy pigment may be a function of the habitat in which the larvae occur rather than an indication of relationships. Use of Larval Characters to Assess Relationships between AGKMS Families.- The AGKMS larvae described in this study possess most of the characters that comprise the distinctive larval form as defined by Johnson (1984), Johnson and Fritzsche (1989) and Leis and van der Lingen (1997): a larval form with strong similarities in body form, generalized head spination, fin development and pigment patterns. The similarities in all four main character groups present in various combinations and degrees of development in our AGKMS larvae are consistent with a close relationship among all five families. However, a comparison of these characters indicates that there is considerable inter-familial variability among our AGKMS larvae and across larvae of other RLA-lO percoid families, and that many of the characters are also present in larvae of non RLA-lO percoid families. In fact, larvae of the non RLA-lO percoid family Pomatomidae have all the characters of the distinctive GKS larval form and are more similar to the AGKMS larval form than are the larvae of the other RLA-lO percoid families (Leis and Rennis, 1983; Johnson, 1984; akiyama, 1988; Leis and Trnski, 1989; Trnski et aI., 1997). Based on larval characters only, one could hypothesize that all RLA-lO percoid families, including also the Dichistiidae (Leis and van der Lingen, 1997), form part of a larger group perhaps including non RLA-lO families such as the 136 BULLETIN OF MARINE SCIENCE. VOL. 60. NO. I. 1997

Pomatomidae. Alternatively, it is also possible that the "distinctive larval form" present in Pomatomus and, to some extent, in some other non RLA-lO families, could have been acquired independently, perhaps related to habitat. Since it is not possible to conclude that any of the attributes of the distinctive larval form constitute synapomorphies, these alternatives could not be tested during this study. Further work is needed to assess the occurrence and homology of the components of the distinctive larval form in other families in order to identify possible out- groups. Information provided in this study suggests that the use of larval characters alone to assesses relationships, both within RLA-lO families as well as across non RLA-lO percoids, is problematical and must be treated with caution. The fact that none of the characters of the distinctive larval form in our AGKMS larvae is exclusive to RLA-lO percoid families implies that larval characters do not by themselves provide convincing evidence of monophyly of these families. Larval AGKMS are generalized and the evidence from the larval morphology is not sufficient to clarify relationships. A different situation pertains with groups such as serranid anthiines and epinephelines, in which specialized larval characters have provided convincing evidence ofmonophyly (Johnson, 1988; Baldwin, 1990; Baldwin and Johnson, 1993).

ACKNOWLEDGMENTS

We wish to acknowledge the following people for their help in providing material and discussion: J. M. Leis (Australian Museum, Sydney), G. D. Johnson (Smithsonian Institution, Washington, D.C.), H. S. Gill (Murdoch University), J. Young (CSIRO, Hobart), and D. Rissik and 1. M. Suthers (Uni- versity of New South Wales, Sydney). We also thank T Trnski (Australian Museum, Sydney) for kindly illustrating the larvae in Figure 2. The editorial comments and suggestions of J. M. Leis, and those of three anonymous reviewers greatly improved the paper and are very much appreciated.

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DATEACCEPTED: March 26, 1996.

ADDRESSES: (F.J.N.) Marine and Freshwater Resources Institute, P.O. Box 114, Queenscliff. Victoria 3225. Australia; (A.G.M.) A WT EnSight, P.O. Box 73, West Ryde, NSW 2114, Australia; (B.D.B) CSIRO Division of Marine Research, GPO Box 1538, Hobart, Tasmania 7001, Australia.