A gastric-brooding asteroid, Smilasterias multipara

The gastric-brooding asterinid sea star, Smilasterias multipara, broods from late August to early November in the shallow sublittoral zone of southeastern Australia. We observed males and females spawning in the laboratory. They shed gametes through gonopores on the sides of the arms. The eggs were orange, about 1.0 mm in diameter, and heavier than seawater. They were externally fertilized by sperm, and placed into the stomach of the female by the tube feet. Twenty-four hours after fertilization, the first cleavage occurred. Cleavage was equal, total, and radial. Development via a non-feeding lecithotrophic brachiolaria was direct, there being no planktrotrophic bipinnaria or brachiolaria larva. Embryos developed, through wrinkled blastula and gastrula stages, into brachiolariae with arms. All of the surfaces of the brachiolaria were covered by cilia. At metamorphosis, a starfish rudiment appeared on the posterior portion of the larval body, while the anterior portion of the larval body was absorbed. Two months after fertilization, metamorphosis was complete. After metamorphosis, juveniles in the stomach grew six pairs of tube feet in each arm. Juveniles, 3 mm in diameter, emerged from the mouth of the mother in early November. Developmental evidence suggests that this asteroid has evolved mechanisms for the protection of larvae and juveniles from gastric digestion.     

The effect of age at metamorphosis on the transition from larval to adult suspension‐feeding of the slipper limpet Crepidula fornicata

Slipper limpets use different ciliary feeding mechanisms as larvae and adults. Veliger larvae of Crepidula fornicata developed part of the adult feeding apparatus, including ctenidial filaments, neck lobe, and radula, before metamorphosis, but ctenidial feeding did not begin until well after loss of the larval feeding apparatus (velum) at metamorphosis. Earlier initiation of ctenidial feeding by individuals that were older larvae when metamorphosis occurred suggests continued development toward ctenidial feeding during delay of metamorphosis. Early juveniles produced a ciliary current through the mantle cavity and moved the radula in a grasping action before they began to capture algal cells on mucous strands or form a food cord. Either early juveniles could not yet form mucous strands or they delayed their production until development of other necessary structures. The neck canal for transporting food from ctenidium to mouth cannot develop before velar loss. In their first feeding, juveniles fed much like the adults except that the neck canal was less developed and the path of the food cord toward the mouth sometimes varied. As suspension feeders, calyptraeids lack the elaborations of foregut that complicate transition to juvenile feeding for many caenogastropods, but a path for the food cord must develop after velar loss. Why individuals can initiate ctenidial feeding sooner when they are older at metamorphosis is not yet known. The juveniles became sedentary soon after metamorphosis and were not observed to feed by scraping the substratum with the radula, in contrast to the first feeding by juveniles of another calyptraeid species, observed by Montiel et al. ( 2005 ).     

Short-term fluctuation in salinity promotes rapid larval development and metamorphosis in Dendraster excentricus

The effect of constant and fluctuating salinity on larval development and metamorphosis of the sand dollar Dendraster excentricus was investigated in the laboratory. Sand dollar larvae at different stages of development were kept either at 32‰ (controls), exposed to constant low salinity (22‰) throughout development, or exposed to fluctuating salinity (i.e. transferring larvae from 32‰ to 22‰ for 7 days then back to 32‰ for the rest of their development). Larvae exposed to constant low salinity were significantly smaller but developed all larval arms at a slower rate than larvae in all other treatments. Larvae exposed to fluctuating salinity recovered and developed significantly longer larval arms and bigger rudiments than larvae kept at constant low salinity. Larvae exposed to fluctuating salinity produced more juveniles than larvae at constant high salinity (32‰), while those at constant low salinity produced few or no juveniles. Four-arm larvae exposed to fluctuating salinity produced significantly more juveniles than six-arm larvae exposed to the same treatment. Transferring competent 8-arm larvae from 31‰ to 15‰ for 2 days then back to 31‰, induced metamorphosis with juvenile production being significantly higher than for those kept at a constant salinity of 20, 25 and 31‰. This study indicates that a short-term decrease in salinity might induce metamorphosis for this species.     

Larval carry‐over effects from ocean acidification persist in the natural environment

An extensive body of work suggests that altered marine carbonate chemistry can negatively influence marine invertebrates, but few studies have examined how effects are moderated and persist in the natural environment. A particularly important question is whether impacts initiated in early life might be exacerbated or attenuated over time in the presence or absence of other stressors in the field. We reared Olympia oyster (Ostrea lurida) larvae in laboratory cultures under control and elevated seawater pCO₂ concentrations, quantified settlement success and size at metamorphosis, then outplanted juveniles to Tomales Bay, California, in the mid intertidal zone where emersion and temperature stress were higher, and in the low intertidal zone where conditions were more benign. We tracked survival and growth of outplanted juveniles for 4 months, halfway to reproductive age. Survival to metamorphosis in the laboratory was strongly affected by larval exposure to elevated pCO₂ conditions. Survival of juvenile outplants was reduced dramatically at mid shore compared to low shore levels regardless of the pCO₂ level that oysters experienced as larvae. However, juveniles that were exposed to elevated pCO₂ as larvae grew less than control individuals, representing a larval carry‐over effect. Although juveniles grew less at mid shore than low shore levels, there was no evidence of an interaction between the larval carry‐over effect and shore level, suggesting little modulation of acidification impacts by emersion or temperature stress. Importantly, the carry‐over effects of larval exposure to ocean acidification remained unabated 4 months later with no evidence of compensatory growth, even under benign conditions. This latter result points to the potential for extended consequences of brief exposures to altered seawater chemistry with potential consequences for population dynamics.     

Spatiotemporal occurrence and feeding habits of tonguefish,Cynoglossus lighti Norman, 1925, larvae in Ariake Bay,Japan

Spatiotemporal occurrence and feeding habits of tonguefish (Cynoglossus lighti Norman, 1925) larvae were investigated in an offshore area (>5 m in depth) of the inner part of Ariake Bay, Japan. All specimens were symmetric, free‐swimming larvae. Although their seasonal abundance and distribution in the study site varied from year to year, spawning started in June and the larval abundance was high in August and September with a wide distribution in the inner part of the bay. Both present and previous study results strongly suggest that larvae may settle primarily in the estuary and near‐shore areas of Ariake Bay after their wide distribution in the offshore area of the bay during the free‐swimming stage. Larvae showed a clear feeding rhythm in which they fed on prey mainly during the daytime. Larvae fed exclusively on copepods, and identified prey were mostly Paracalanidae (mainly Parvocalanus crassirostris), Microsetella norvegica, and Oithona davisae. Pre‐metamorphosis larvae fed primarily on Paracalanidae and O. davisae, whereas O. davisae formed a smaller proportion of the early‐metamorphosis diet. In early metamorphoses, larvae fed preferentially on Paracalanidae and M. norvegica.     

Effects of temperature on survival, development, growth and feeding of larvae of Yellowtail clownfish Amphiprion clarkii (Pisces: Perciformes)

To understand the physiological and ecological responses of marine fishes to the change of water temperature, newly-hatched larvae of Yellowtail clownfish Amphiprion clarkii were reared in captivity at water temperatures of 23, 26 and 29 °C till they completed the metamorphosis to juvenile phase, and larval survival, development, growth and feeding were evaluated during the experimental period. The results showed that water temperature influenced the physiological performance of larvae of A. clarkii significantly. The survival and growth rates of larvae of A. clarkii increased significantly with the increase of water temperature from 23 to 29 °C (P < 0.05). Water temperature also influenced larval development of A. clarkii significantly and larvae reared at 23 °C took longer time for post-larval development and metamorphosis compared to 26 and 29 °C (P < 0.05). Total length and body weight for post-larval development and metamorphosis decreased with the increase of water temperature from 23 to 29 °C (P < 0.05). Q₁₀ in developmental rate was higher than in daily growth rate at the same rearing temperature, indicating that at water temperature had greater influence on larval development than on growth. Water temperature also influenced larval feeding of A. clarkii significantly with feed ration (FR) and feed conversion efficiency (FCE) increased with the increase of water temperature from 23 to 29 °C (P < 0.05). There was a positive correlation between FR and specific growth rate (SGR) (P < 0.05) but not between FCE and SGR (P > 0.05), indicating that FR influenced growth rate significantly in larvae of A. clarkii. This study demonstrated that the physiological responses of larvae of A. clarkii to the change of water temperature and confirmed that water temperature influenced larval survival, development, growth and feeding significantly. This study suggests that the decline of larval survival and growth rates, extension of pelagic larval duration and reduction of larval feeding at lower temperature have ecological impacts on larval dispersal and metamorphosis, juvenile settlement and population replenishment in A. clarkii in the wild.     

Effects of egg size reduction and larval feeding on juvenile quality for a species with facultative-feeding development

In free-spawning marine invertebrates, larval development typically proceeds by one of two modes: planktotrophy (obligate larval feeding) from small eggs or lecithotrophy (obligate non-feeding) from relatively large eggs. In a rare third developmental mode, facultative planktotrophy, larvae can feed, but do not require particulate food to complete metamorphosis. Facultative planktotrophy is thought to be an intermediate condition that results from an evolutionary increase in energy content in the small eggs of a planktotrophic ancestor. We tested whether an experimental reduction in egg size is sufficient to restore obligate planktotrophy from facultative planktotrophy and whether the two sources of larval nutrition (feeding and energy in the egg) differentially influence larval survival and juvenile quality. We predicted, based on its large egg size, that a reduction in egg size in the echinoid echinoderm Clypeaster rosaceus would affect juvenile size but not time to metamorphosis. We reduced the effective size of whole (W) zygotes by separating blastomeres at the two- or four-cell stages to create half- (H) or quarter-size (Q) “zygotes” and reared larvae to metamorphosis, both with and without particulate food. Larvae metamorphosed at approximately the same time regardless of food or egg size treatment. In contrast, juveniles that developed from W zygotes were significantly larger, had higher organic content and had longer and more numerous spines than juveniles from H or Q zygotes. Larvae from W, H and Q zygotes were able to reach metamorphosis without feeding, suggesting that the evolution of facultative planktotrophy in C. rosaceus was accompanied by more than a simple increase in egg size. In addition, our results suggest that resources lost by halving egg size have a larger effect on larval survival and juvenile quality than those lost by withholding particulate food.     

Testing the evolutionary constraints of metamorphosis: The ontogeny of head shape in Triturus newts

In vertebrates with complex, biphasic, life cycles, larvae have a distinct morphology and ecological preferences compared to metamorphosed juveniles and adults. In amphibians, abrupt and rapid metamorphic changes transform aquatic larvae to terrestrial juveniles. The main aim of this study is to test whether, relative to larval stages, metamorphosis (1) resets the pattern of variation between ontogenetic stages and species, (2) constrains intraspecific morphological variability, and (3) similar to the “hour‐glass” model reduces morphological disparity. We explore postembryonic ontogenetic trajectories of head shape (from hatching to completed metamorphosis) of two well‐defined, morphologically distinct Triturus newts species and their F1 hybrids. Variation in head shape is quantified and compared on two levels: dynamic (across ontogenetic stages) and static (at a particular stage). Our results show that the ontogenetic trajectories diverge early during development and continue to diverge throughout larval stages and metamorphosis. The high within‐group variance and the largest disparity level (between‐group variance) characterize the metamorphosed stage. Hence, our results indicate that metamorphosis does not canalize head shape variation generated during larval development and that metamorphosed phenotype is not more constrained relative to larval ones. Therefore, metamorphosis cannot be regarded as a developmental constraint, at least not for salamander head shape.     

Thyroid hormones determine developmental mode in sand dollars (Echinodermata: Echinoidea)

Evolutionary transitions in larval nutritional mode have occurred on numerous occasions independently in many marine invertebrate phyla. Although the evolutionary transition from feeding to nonfeeding development has received considerable attention through both experimental and theoretical studies, mechanisms underlying the change in life history remain poorly understood. Facultative feeding larvae (larvae that can feed but will complete metamorphosis without food) presumably represent an intermediate developmental mode between obligate feeding and nonfeeding. Here we show that an obligatorily feeding larva can be transformed into a facultative feeding larva when exposed to the thyroid hormone thyroxine. We report that larvae of the subtropical sand dollar Leodia sexiesperforata (Echinodermata: Echinoidea) completed metamorphosis without exogenous food when treated with thyroxine, whereas the starved controls (no thyroxine added) did not. Leodia sexiesperforata juveniles from the thyroxine treatment were viable after metamorphosis but were significantly smaller and contained less energy than sibling juveniles reared with exogenous food. In a second starvation experiment, using an L. sexiesperforata female whose eggs were substantially larger than in the first experiment (202+/-5 vs. 187+/-5 microm), a small percentage of starved L. sexiesperforata larvae completed metamorphosis in the absence of food. Still, thyroxine-treated larvae in this experiment completed metamorphosis faster and in much higher numbers than in the starved controls. Furthermore, starved larvae of the sand dollar Mellita tenuis, which developed from much smaller eggs (100+/-2 microm), did not complete metamorphosis either with or without excess thyroxine. Based on these data, and from recent experiments with other echinoids, we hypothesize that thyroxine plays a major role in echinoderm metamorphosis and the evolution of life history transitions in this group. We discuss our results in the context of current life history models for marine invertebrates, emphasizing the role of egg size, juvenile size, and endogenous hormone production for the evolution of nonfeeding larval development.     

Gregariousness does not vary with geography,developmental stage,or group relatedness in feeding redheaded pine sawfly larvae

Aggregations are widespread across the animal kingdom, yet the underlying proximate and ultimate causes are still largely unknown. An ideal system to investigate this simple, social behavior is the pine sawfly genus Neodiprion, which is experimentally tractable and exhibits interspecific variation in larval gregariousness. To assess intraspecific variation in this trait, we characterized aggregative tendency within a single widespread species, the redheaded pine sawfly (N. lecontei). To do so, we developed a quantitative assay in which we measured interindividual distances over a 90‐min video. This assay revealed minimal behavioral differences: (1) between early‐feeding and late‐feeding larval instars, (2) among larvae derived from different latitudes, and (3) between groups composed of kin and those composed of nonkin. Together, these results suggest that, during the larval feeding period, the benefits individuals derive from aggregating outweigh the costs and that this cost‐to‐benefit ratio does not vary dramatically across space (geography) or ontogeny (developmental stage). In contrast to the feeding larvae, our assay revealed a striking reduction in gregariousness following the final larval molt in N. lecontei. We also found some intriguing interspecific variation: While N. lecontei and N. maurus feeding larvae exhibit significant aggregative tendencies, feeding N. compar larvae do not aggregate at all. These results set the stage for future work investigating the proximate and ultimate mechanisms underlying developmental and interspecific variation in larval gregariousness across Neodiprion.     

Low salinity stress experienced by larvae does not affect post-metamorphic growth or survival in three calyptraeid gastropods

Marine larvae that experience some sub-lethal stresses can show effects from those stresses after metamorphosis, even when they seem to recover from those stresses before metamorphosis. In this study we investigated the short and long-term effects of exposing the larvae of three calyptraeid gastropods (Crepidula fornicata, Crepidula onyx, and Crepipatella fecunda) to temporary reductions in salinity. Larvae of all three species showed slower larval growth rates, longer time to metamorphic competence, and substantial mortality after being stressed in seawater at salinities of 10, 15, and 20 for less than 48 h. Larval tolerance to low salinities varied widely within and among species, but longer stresses at lower salinities were generally more harmful to larvae. However, larvae in nearly all experiments that were able to metamorphose survived and grew normally as juveniles; there were no documented “latent effects.” For all three species, starving larvae in full-strength seawater was not as harmful as exposing larvae to low salinity stress, indicating that detrimental effects on larvae were caused by the salinity stress per se, rather than by an indirect effect of salinity stress on feeding. C. fornicata that were stressed with low salinity as juveniles were more tolerant of the stress than larvae: all stressed juveniles lived and showed reduced growth rates for no more than 3 days. Our data suggest that even though reduced salinity is clearly stressful to the larvae of these 3 gastropod species, metamorphosis seems to generally provide individuals with a fresh start.     

Growth and morphological development of laboratory-reared larval and juvenile three-spot gourami <Emphasis Type="Italic">Trichogaster trichopterus</Emphasis>

Morphological development, including the body proportions, fins, pigmentation and labyrinth organ, in laboratory-hatched larval and juvenile three-spot gourami Trichogaster trichopterus was described. In addition, some wild larval and juvenile specimens were observed for comparison. Body lengths of larvae and juveniles were 2.5 ± 0.1 mm just after hatching (day 0) and 9.2 ± 1.4 mm on day 22, reaching 20.4 ± 5.0 mm on day 40. Aggregate fin ray numbers attained their full complements in juveniles >11.9 mm BL. Preflexion larvae started feeding on day 3 following upper and lower jaw formation, the yolk being completely absorbed by day 11. Subsequently, oblong conical teeth appeared in postflexion larvae >6.4 mm BL (day 13). Melanophores on the body increased with growth, and a large spot started forming at the caudal margin of the body in flexion postlarvae >6.7 mm BL, followed by a second large spot positioned posteriorly on the midline in postflexion larvae >8.6 mm BL. The labyrinth organ differentiated in postflexion larvae >7.9 mm BL (day 19). For eye diameter and the first soft fin ray of pelvic fin length, the proportions in laboratory-reared specimens were smaller than those in wild specimens in 18.5–24.5 mm BL. The pigmentation pattern of laboratory-reared fish did not distinctively differ from that in the wild ones. Comparisons with larval and juvenile morphology of a congener T. pectoralis revealed several distinct differences, particularly in the numbers of myomeres, pigmentations and the proportional length of the first soft fin ray of the pelvic fin.     

Development of the starfish <Emphasis Type="Italic">Asterias amurensis</Emphasis> under laboratory conditions

Under laboratory conditions the development of the starfish Asterias amurensis Lütken from Vostok Bay (Sea of Japan) was studied at 14 and 17°C. At 14°C and a salinity of 31.6–32.6, ciliated coeloblastulae hatched from egg envelopes 19 h after fertilization. At this temperature the development proceeded slowly and stopped at the stage of bipinnaria. At 17°C and normal salinity of seawater, the development of A. amurensis was successful. The swimming blastula appeared in 14 h. It took 30.5 h for the embryos to reach the gastrula stage. The larvae began swimming in a horizontal position with the apical tip ahead. The dipleurula appeared at 60 h. These larvae began feeding. At 71 h after the beginning of development, the early bipinnaria has developed. In the larva, the edged ciliated band, the preoral plate, and the anal plate were already formed. At the age of 4.2 days, the larvae reached the stage of bipinnaria and the brachiolaria stage developed by 26–28 days after fertilization. The larvae had three identical brachiolar arms with attachment papillae on their tips and an attachment disk. In 37–44 days (at 17°C) the pelagic phase of A. amurensis development was completed by the attachment of larvae to the bottom plates and termination of metamorphosis. Most likely, the specificity to a substrate is not expressed in the brachiolaria of A. amurensis. They can settle on almost any hard substrate which is coated with a bacterial film. The newly settled juvenile starfish had five well-developed arms and moved using their ambulacral podia.Original Russian Text Copyright © 2005 by Biologiya Morya, Kashenko.     

Morphogenesis and phenotypic divergence in two developmental morphs of Streblospio benedicti (Annelida,Spionidae)

Abstract. The morphology of marine invertebrate larvae is strongly correlated with egg size and larval feeding mode. Planktotrophic larvae typically have suites of morphological traits that support a planktonic, feeding life style, while lecithotrophic larvae often have larger, yolkier bodies, and in some cases, a reduced expression of larval traits. Poecilogonous species provide interesting cases for the analysis of early morphogenesis, as two morphs of larvae are produced by a single species. We compared morphogenesis in planktotrophic and lecithotrophic morphs of the poecilogonous annelid Streblospio benedicti from the trochophore stage through metamorphosis, using observations of individuals that were observed alive, with scanning electron microscopy, or in serial sections. Offspring of alternate developmental morphs of this species are well known to have divergent morphologies in terms of size, yolk content, and the presence of larval bristles. We found that some phenotypic differences between morphs occur as traits that are present in only one morph (e.g., larval bristles, bacillary cells on the prostomium and pygidium), but that much of the phenotypic divergence is based on heterochronic changes in the differentiation of shared traits (e.g., gut and coelom). Tissue and organ development are compared in both morphs in terms of their structure and ontogenetic change throughout early development and metamorphosis.     

Are the anticipatory pathways in lecithotrophic larvae that delay metamorphosis adaptations? (A review)

During anticipatory development in lecithotrophic larvae that delay metamorphosis, the growth and differentiation of features of the adult action system continue to develop at a slow pace even though they do not become functional. After metamorphosis occurs, the larger size and advanced development of these components may allow juveniles to initially grow at a faster rate than they normally would. Anticipatory development has been demonstrated in archeogastropods, some solitary ascidians and a hydrozoan. In the gastropod Haliotis and the hydrozoan Phialidium anticipatory development increases the initial growth rate of juveniles. In Haliotis and ascidians all of the larvae of a given female that live long enough exhibit anticipatory development. In Phialidium, the ability of a given female to produce larvae that can exhibit anticipatory development is a maternal polymorphic character. In Haliotis and solitary ascidians that exhibit anticipatory development, it appears to be a slower version of the rapid developmental changes that occur in parts of the adult action system at metamorphosis. In Phialidium, developmental changes in relative sizes of the different presumptive regions of the polyp are slowly altered prior to and independently of metamorphosis. Anticipatory development is not linked to the decrease in the size or nutrient reserves of older larvae but to the length of their larval period. From an evolutionary perspective, the mechanisms that operate during anticipatory development are probably of adaptive significance for lecithotrophic larvae of species that spend variable amounts of time in the water column because of a patchy distribution of appropriate settlement cues. The developmental mechanisms that underlie anticipatory development may have been used during the transition from lecithotrophy to planktotrophy.     

Olfactory cues used in host‐habitat location and host location by the parasitoid Cotesia urabae

There is a growing body of evidence that many hymenopteran parasitoids make use of olfaction as the primary mechanism to detect and locate hosts. In this study, a series of bioassays was conducted to investigate the orientation behaviour of the gum leaf skeletonizer larval parasitoid Cotesia urabae Austin & Allen (Hymenoptera: Braconidae) in both Y‐tube and four‐arm olfactometers. In a Y‐tube olfactometer, male C. urabae were attracted only to virgin conspecific females. Host‐plant leaves, damaged leaves, host larvae, and host larvae feeding on leaves were highly attractive to female C. urabae, whereas host frass and conspecific males were not. The multiple‐comparison bioassay conducted in a four‐arm olfactometer clearly indicates that C. urabae females were significantly more attracted to the host Uraba lugens Walker (Lepidoptera: Nolidae) larvae feeding on Eucalyptus fastigata H Deane & Maiden (Myrtaceae) leaves than to any other of the odour sources tested. The results of this study show that C. urabae individuals responded to chemical cues specific to the host plant and target host insect, and support hypotheses that unreliable cues are not utilized for host location by specific natural enemies.     

The effects of nurse eggs and sibling interactions on the larval development of the poecilogonous annelid Boccardia proboscidea (Spionidae)

In poecilogony, different types of larvae are produced within the same species. Previous studies have suggested maternal control of the production of larval types; however, it is not clear which factors or mechanisms generate contrasting developmental patterns among siblings. The spionid polychaete Boccardia proboscidea produces within the same capsule adelphophagic larvae that eat nurse eggs and siblings and complete all or most of their development inside the capsule (Type A larvae), and larvae with little growth until they hatch as planktotrophic larvae (Type B larvae). In this study, we manipulated capsule content to explore the factors determining larval type in B. proboscidea and the role of extra‐embryonic maternal nutrition and sib–sib interaction in the developmental fate of offspring. When early larval stages were grown individually in vitro, with nurse eggs as the only food source, some of them remained small, while others continue developing into larger pre‐competent larvae by feeding on nurse eggs. This suggests that larval types in B. proboscidea are determined very early in development and are not solely the product of sib–sib interaction inside the capsule. However, our data also suggest that hatching size variability within larval types of a clutch depends on nurse egg availability. Type B larvae grew normally to metamorphosis when phytoplankton was available, but suffered high rates of cannibalism by Type A larvae. These results are consistent with the hypothesis that individual larval fates are determined very early in development and that once their fate is determined, hatching size and intracapsular survival are affected by maternal food provisioning and sibling interaction.     

Growth and morphological development of laboratory-reared larval and juvenile Pangasianodon hypophthalmus

The morphological development, including the fins, body proportions and pigmentation, of laboratory-reared larval and juvenile Pangasianodon hypophthalmus was described and their behavioral features were observed under rearing conditions. Body lengths (BL) of larvae and juveniles were 3.0 ± 0.2 (mean ± SD) mm just after hatching, and 12.9 ± 1.1 mm on day 13, reaching 23.4 ± 1.8 mm on day 25 after hatching. Aggregate fin ray numbers (for caudal fin, principal soft ray number) attained their full complements in specimens larger than 12.8 mm BL. Notochord flexion began in yolksac larvae on day 0 (10.5 h after hatching), with teeth buds and barbels appearing with jaw formation in yolksac flexion larvae on day 1. Melanophores on the body increased with growth, with a broad vertical band forming on the lateral line and an oblique band extending from above the pectoral fin base towards the forepart of the anal fin during the postflexion larval and juvenile stages. Body proportions became relatively constant in juveniles, except for maxillary barbel length (MBL), which continued to decrease. Yolksac flexion larvae started feeding on day 2 with the onset of intense cannibalism. Yolks were completely absorbed by day 3, and cannibalism ended by day 6. Subsequently, fish displayed a schooling behavior with growth, preferring relatively dark areas during the juvenile stage.