Osteological development of the vertebral column and of the fins in Diplodus sargus (Teleostei: Perciformes: Sparidae)

The osteological development of vertebral column and fins in white sea bream, Diplodus sargus (Linnaeus, 1758) was studied. Vertebral ontogeny started at 5.7 mm total length (TL, post-fixation), with the formation of the first cartilaginous neural and haemal arches, and was completed by the full attainment of dorsal ribs (20.8 mm TL). The formation of vertebral centra occurred between 7.3 and 9.4 mm TL, but was fully completed only after the development of post- and pre-zygapophyses (26.0 mm TL). Pectoral supports were the first fin elements that started to develop (3.8 mm TL), followed by those of the caudal fin (5.2 mm TL), dorsal and anal fins (6.7 mm TL), and then by those of the pelvic fin (9.5 mm TL). The caudal fin of D. sargus was the first to develop fin rays and attain the full count of lepidotrichia (5.6-7.8 mm TL), but the last to attain the full count of dermatotrichia (7.9-16.0 mm TL). The next fins starting to present rays were the pectoral (6.5 mm TL) and the dorsal and anal fins (7.5 mm TL), while the pelvic fins were the last (11.5 mm TL). Following the caudal lepidotrichia (7.8 mm TL), the anal (10.5 mm TL), dorsal (12.0 mm TL), pectoral (11.4 mm TL) and pelvic fins (12.5 mm TL) were the next with fully completed ray counts. Finally, fin meristics were fully developed with the caudal dermatotrichia. The results are discussed in comparison with the fin and vertebral ontogeny of other teleosts.     

Development of the skull in Dentex dentex (Osteichthyes: Sparidae)

The development of the skull skeleton in common dentex, Dentex dentex (Linnaeus, 1758), was studied under extensive rearing conditions. Skull ontogeny started after hatching and went through three stages: (a) the yolk-sac (during the phase of no growth) and the early larval stage (up to 6.9 mm in vivo total length, TL) in which the majority of the skull elements formed; (b) the late larval stage (up to 11.3 mm TL) in which the basisphenoid, infraorbital 1, parethmoids, nasals, ethmoids, pleurosphenoids, opisthotics and orbitosphenoids developed; and (c) the metamorphosis stage (up to 24.0 mm TL) during which the remaining infraorbitals formed, the cartilaginous areas of the neu- roskull regressed and the sclerotics started to ossify. Teeth first appeared at 5.2 mm TL, and gill filaments formed between 5.7 and 6.9 mm TL. The results are discussed in view of the ontogeny of functions and in comparison with the skull ontogeny of other teleosts. Received: 1 July 1998 / Accepted: 1 September 1999     

Ontogeny and allometric plasticity of Dentex dentex (Osteichthyes: Sparidae) in rearing conditions

The ontogeny of Dentex dentex (Linnaeus, 1758) was studied comparatively under extensive and semi-extensive rearing conditions. The exotrophic phase was divided into three distinct developmental stages (a) larval, from feeding onset up to 10 mm total length (TL); (b) metamorphosis, from 10 mm TL up to 24 mm TL; and (c) further juveniles. The majority of all ontogenic changes was expressed during the larval stage and integrated during the metamorphosis stage. The juvenile stage was characterized by developmental stability. The normal development of D. dentex was seriously affected by the rearing conditions applied during the early exotrophic phase. The two differently reared populations followed similar developmental patterns of general morphology, pigmentation and behaviour, but they presented high developmental plasticity where allometric growth was concerned. Rearing conditions influenced both the boundaries of the sequential stages (in 50% of the morphometric ratios) and the allometry coefficient of each morphometric character in the various developmental stages (in 80% of the characters). The results are discussed with respect to the ontogeny of the functional morphology under both ecological and aquaculture considerations. Received: 1 August 1998 / Accepted: 1 September 1999     

Asymptotic growth trajectories of larval sardine (Sardinops melanostictus ) in the coastal waters off western Japan

Growth trajectories of individual larvae of Japanese sardine, Sardinops melanostictus, caught in the coastal waters off western Japan were back-calculated from the first feeding stage up to date of capture (approximate size of 20 to 35 mm total length; TL) based on individually determined allometric relationships between otolith daily ring radii and fish total lengths. The larvae in January-, February-, and March-hatched cohorts in the coastal waters grew faster and more uniformly than those in the oceanic waters offshore of the Kuroshio current. Growth trajectories of the three hatch-month cohorts were similar and could be expressed by the Gompertz model. The inflection points of the growth curves were reached at 9 to 11 d after hatching, when larvae were 10.8 to 11.8 mm TL. Maximum growth rates at these points were 0.80 to 0.85 mm d⁻¹. Growth rates gradually declined after the inflection points, and larval TLs converged into the infinite length of 29 to 32 mm, the sizes at which metamorphosis from larvae to juveniles is initiated. This asymptotic growth pattern in the larval stage resulted in the narrow ranges in TLs in spite of the wide range of ages of the larvae caught by boat seiners in the coastal waters. Slow growth and therefore long duration of the metamorphosing stage could be influential in determining the cumulative total mortality in the early life stages of the Japanese sardine. Received: 14 July 1996 / Accepted: 20 August 1996     

Ontogenetic variation in metabolism, biochemical composition and energy content during the early life stages of Farfantepenaeus paulensis (Crustacea: Decapoda: Penaeidae)

Dry weight (DW), oxygen consumption, ammonia-N excretion, proximate biochemical composition (total protein, carbohydrate, lipid, water and ash), and energy content (estimated from biochemical composition and by wet combustion) were determined in early developmental stages of cultured Farfantepenaeus paulensis. Pooled samples from embryonic, larval and postlarval stages (at 26 ± 1 °C and 34 ± 1‰) were used for measurements. The study focused on physiological and biochemical processes during transitional periods of ontogeny, such as hatching, lecithotrophic and planktotrophic stages, metamorphosis, and the attainment of a benthic existence in postlarva. DW showed higher increment between protozoea I (PZ I) and mysis I (M I) than in the next mysid and postlarval stages. Individual rates of oxygen consumption and ammonia-N excretion increased, while weight-specific rates presented significant reduction throughout development. Higher weight-specific oxygen consumption was registered in nauplius III (N III) and PZ I, following a decrease in subsequent stages. Postlarval stages PL V–VI and PL X–XII exhibited the lowest values among the stages studied. Weight-specific excretion was high in N III and protozoeal stages, with maximum values in PZ II, while the following stages were marked by lower rates. O:N ratios indicated higher protein catabolism in the stages between egg and M I and a shift to more lipid utilization close to metamorphosis. Water content was higher in the protozoeal stages and decreased afterwards. Higher percentages of protein, lipid and carbohydrate (%DW) were observed in egg and nauplius stages. Protein and lipid decreased from the egg through the naupliar and protozoeal stages, rising again in mysis stages. Lipid content (%DW) decreased in PL V–VI and PL X–XII. Lipid:protein ratios showed an increase of the importance of lipid between PZ III and M II. Carbohydrates represented a minor fraction of body composition, and ash percentages increased from egg to a maximum in PZ II, decreasing in subsequent stages. Energy content determined by wet combustion or calculated by energy equivalents presented the same trend throughout development, varying similarly to protein. Protein was the main energy contributor to body energy in all stages, while the importance of lipid was higher in egg and early naupliar stages. Trends observed in metabolic rates and body composition may be associated to morphological and behavioral changes during the early stages of penaeid development, such as the transition from herbivory to omnivory, and the adoption of a benthic existence. Different ontogenetic energy strategies contribute to succeed through such diverse type of development. Received: 4 July 2000 / Accepted: 6 December 2000     

Gill morphogenesis in the oyster Ostrea chilensis

Despite the importance of the gills in the acquisition of food by suspension-feeding bivalve mollusks, there is almost no information on gill organogenesis. By means of a series of stereoscan electron micrographs, this paper describes gill development in the Chilean oyster, Ostrea chilensis, from the brooded larval stages to 1-month-old spat. A single gill rudiment was observed on each side of the mantle at a shell length of 320 μm, and the rudiments increased in number and size until the end of the brooding period. During metamorphosis the gill filaments increased in number from 5 or 6 to between 7 and 9. The loss of the velum and the absence of functional gill filaments during metamorphosis are consistent with previous observations of weight loss during this critical period of the life history, because the newly settled juvenile lacks the ability to remove particles from suspension. The end of metamorphosis (100% of spat with dissoconch edge) was reached 36 h after larval settlement, when the gill filaments began to grow cilia, which increased in density and differentiated as the spat developed and acquired the capability of suspension-feeding, accounting for the increase in body weight previously recorded during this stage. The larval rudiments gave rise to the inner demibranchs. The outer demibranchs were observed 10 days after settlement, located between the inner demibranch and the mantle. In 1-month-old spat, the gill did not show differentiation between primary and secondary filaments, indicating that the heterorhabdic condition characteristic of adult oysters had yet to be attained. Received: 11 December 1998 / Accepted: 21 August 2000     

Role of biochemical energy reserves in the metamorphosis and early juvenile development of the oyster Ostrea chilensis

Metamorphosis in the Chilean oyster Ostrea chilensis was complete 36 h after release of the larvae, when 100% of the individuals showed edge growth of the dissoconch. The size of the larval shell did not change during metamorphosis, although the total dry weight of the larva decreased considerably. During this period, when the gill ciliature was undeveloped and the oyster therefore unable to feed, energy demands were met by biochemical reserves retained from the larval phase. Proteins contributed the largest quantity of energy to the metamorphosing oyster, 69.3% of the total expended, whereas lipids supplied 24.3% and carbohydrates only 6.4%. The process of metamorphosis consumed 64.5% of the energy reserves held by the pediveliger at the time of release. When metamorphosis was complete, growth began and tissue reserves were replenished, protein and carbohydrate accumulating rapidly early in the juvenile stage. Received: 26 December 1997 / Accepted: 8 July 1998     

Role of protein in larval swimming and metamorphosis of Bugula neritina (Bryozoa: Cheilostomatida)

Larvae of the marine cheilostomatid bryozoan Bugula neritina (L.) were prevented from settling for 1, 4 and 8 h by mechanical agitation, following which settlement and metamorphosis success were examined. Settlement rates were significantly affected by swimming time, which decreased from 100% after 2 h to 93.7 ± 4.3% after 8 h. Similarly, metamorphosis to the feeding ancestrula was significantly impaired following a swimming time of 8 h, declining from 93.7 ± 4.3% after 1 h to 65.9 ± 7.0% after 8 h. The resultant colonies grew well for the first 3 wk, following which time, growth patterns became erratic. Growth rate was in all cases highly variable, and did not correlate with enforced swimming times. Larval protein composition was examined after 1, 4 and 8 h swimming time, and post-larval composition 1, 2, 5, 24 and 48 h after settlement using sodium-dodecyl-sulphate polyacrylamide-gel electrophoresis (SDS-PAGE). Individual protein content was measured using a densitometer. Larvae did not consume protein during swimming, however a protein measuring 170 kdaltons was consumed during metamorphosis. These results are discussed in the context of larval settlement and energetics. Received: 19 July 1998 / Accepted: 3 December 1998     

Facultative lecithotrophy during larval development of the burrowing shrimp Callianassa tyrrhena (Decapoda: Callianassidae)

Survival, developmental and consumption rate (Artemia nauplii ingested per day) as well as predation efficiency (ingested per available Artemia nauplii) were studied during the larval development of the shallow-water burrowing thalassinid Callianassa tyrrhena (Petagna, 1792), which exhibits an abbreviated type of development with only two zoeal stages and a megalopa. The larvae, hatched from berried females from S. Euboikos Bay (Aegean Sea, Greece), were reared at 10 temperature–food density combinations (19 and 24 °C; 0, 2, 4, 8 and 16 Artemia nauplii d⁻¹). Enhanced starvation resistance was evident: 92 and 58% of starved zoeas I molted to zoea II, while metamorphosis to megalopa was achieved by 76 and 42% of the hatched zoeas at 19 and 24 °C, respectively. The duration of both zoeal stages was affected by temperature, food density and their interaction. Nevertheless, starvation showed different effects at the two temperatures: compared to the fed shrimp, the starved zoeae exhibited accelerated development at 19 °C (8.4 d) but delayed metamorphosis at 24 °C (5.9 d). On the other hand, both zoeal stages were able to consume food at an increased rate as food density and temperature increased. Predation efficiency also increased with temperature, but never exceeded 0.6. Facultative lecithotrophy, more pronounced during the first zoeal stage of C.tyrrhena, can be regarded as an adaptation of a species whose larvae can respond physiologically to the different temperature–food density combinations encountered in the wide geographical range of their natural habitat. Received: 28 February 1998 / Accepted: 21 October 1998     

Timing of metamorphosis and larval segregation of the Atlantic eels Anguilla rostrata and A. anguilla, as revealed by otolith microstructure and microchemistry

Otolith microstructure and microchemistry were examined in juveniles of American (Anguilla rostrata) and European (A. anguilla) eels. Otolith increment width markedly increased from age 132 to 191 d (156 ± 18.9 d; mean ± SD) in A. rostrata and 163 to 235 d (198 ± 27.4 d; mean ± SD) in A. anguilla, both of which were coincident with drastic decreases in otolith Sr:Ca ratios, suggesting that metamorphosis from leptocephalus to glass eel began at those ages in each species. The duration of metamorphosis was estimated to be 18 to 52 d from otolith microstructure, for both species studied. Ages at recruitment were 171 to 252 d (206 ± 22.3 d; mean ± SD) in A. rostrata and 220 to 281 d (249 ± 22.6 d; mean ± SD) in A. anguilla. In these two species, positive linear relationships were found in ages between the beginning of metamorphosis and recruitment, suggesting that early metamorphosing larvae recruited at younger ages. Duration of the leptocephalus stage to recruitment in A. anguilla was about 40 d longer than that in A. rostrata. The geographical segregation between the two species in the Atlantic Ocean seems to be involved in the differences in the duration of the leptocephalus stage (age at metamorphosis). Received: 8 November 1999 / Accepted: 8 May 2000     

Spawning and larval development of the black turban snail Tegula funebralis (Prosobranchia: Trochidae)

An understanding of spawning and larval development can be fundamental to interpreting the abundance, distribution, and population structure of marine invertebrate taxa. Tegula funebralis (A. Adams, 1855), the black turban snail, has been the focus of numerous ecological studies on the Pacific coast of North America. To date, there are only conflicting and anecdotal reports of spawning, and there is no information on larval or juvenile development for this conspicuous and abundant species. On 19 September 1995, two individuals of T. funebralis were observed free-spawning gametes into seawater in tanks at the Oregon Institute of Marine Biology. Embryos and larvae were subsequently reared to metamorphosis and beyond. Development was pelagic and similar to development described for other trochids, and larvae were observed not to feed at any stage. Larvae began to metamorphose at 5.7 to 6.7 d and settled at 260 μm shell length. Juveniles grew ≃ 10 μm in shell length per day and appeared to feed on detritus. Juveniles lacked some adult diagnostic shell characters, including two columellar nodes and a closed umbilicus. In the field, small (<3 mm) juveniles occurred in the adult habitat on all sampling dates between October and March. Small juveniles were found only under rocks and were most abundant under rocks partially buried in coarse sand, suggesting that juveniles may utilize a specific microhabitat within the adult T. funebralis habitat. Received: 7 October 1996 / Accepted: 17 October 1996     

Genetic heterogeneity among adult and recruit red sea urchins, Strongylocentrotus franciscanus

Allozyme electrophoresis was used to characterize genetic variation within and among natural populations of the red sea urchin Strongylocentrotus franciscanus. In 1995 to 1996, adult urchins were sampled from twelve geographically separated populations, seven from northern California and five from southern California (including Santa Rosa Island). Significant population heterogeneity in allelic frequencies was observed at five of six polymorphic loci. No geographic pattern of differentiation was evident; neighboring populations were often more genetically differentiated than distant populations. Northern and southern populations were not consistently distinguishable at any of the six loci. In order to assess within-population genetic variation and patterns of recruitment, large samples were collected from several northern California populations in 1996 and 1997, and were divided into three size classes, roughly representing large adults (>60 mm), medium-sized individuals (31 to 60 mm, “subadults”) and individuals <2 yr of age (≤30 mm test diam, referred to as “recruits”). Comparisons of allelic counts revealed significant spatial and temporal differentiation among size-stratified population samples. Recruit samples differed significantly from adult samples collected at the same locale, and showed extensive between-year variation. Genetic differentiation among recruit samples was much higher in 1997 than in 1996. Between-year differences within populations were always greater for recruits than for adults. Potential explanations for the differentiation of recruit samples include pre- and post-settlement natural selection and high interfamily variance in reproductive success or “sweepstakes” recruitment. Unless recruit differentiation can be attributed to an improbable combination of strong and spatially diverse selection, such differentiation across northern California populations indicates that the larval pool is not well mixed geographically (even on spatial scales <20 km), despite long planktonic larval duration. Received: 6 July 1999 / Accepted: 25 January 2000     

Growth, carbon, nitrogen and caloric content of Solea senegalensis (Pisces: Soleidae) from egg fertilization to metamorphosis

Eggs and larvae of the Senegal sole, Solea senegalensis Kaup, were reared from fertilization until the end of metamorphosis, which occurs by Day 17 after hatching at 19.5 °C. Changes in energy content and biomass quality were studied in terms of dry weight and of carbon, nitrogen and energy content. S. senegalensis spawned eggs of about 1 mm diameter which hatched 38 h after fertilization. Average dry weight of individual eggs was 46 μg, the chorion accounting for about 18% of total dry weight. Gross energy of recently fertilized sole eggs was approximately 1 J egg⁻¹. From fertilization to hatching, eggs lost 8% of their total energy (chorion not included). After hatching, larvae lost 14% of their initial energy until the start of feeding which occurred about 48 h afterwards. The principal components catabolized during embryogenesis were carbon-rich compounds that decreased by 26%, while nitrogen-rich compounds decreased by only 10% and were practically unaltered from hatching to the start of feeding. Feeding larvae displayed constant growth during the period studied (specific growth rate on a dry weight basis was 0.26 d⁻¹). The relative proportion of carbon and nitrogen content revealed an accumulation of high energy compounds in the days before metamorphosis. By Day 14, the energy content reached values similar to those of recently hatched embryos, but decreased again during metamorphosis. Received: 10 June 1998 / Accepted: 28 January 1999     

Temperature-induced ontogenetic plasticity in sea bass (Dicentrarchus labrax)

We studied the ontogeny of Dicentrarchus labrax comparatively under constant rearing temperatures of 13, 15 and 20°C. At hatching, yolk-sac larval morphometry differed significantly between the temperatures and especially between the two extremes, while at the end of the yolk-sac larval stage, it mainly differed between the two lower temperature regimes and that of 20°C. Compared with the two lower temperature conditions, at 20°C D. labrax presented a significant ontogenetic acceleration which was morphologically expressed either as a significantly smaller total length (TL) at feeding onset, notochord flexion and fin differentiation, or as shifts of the allometric inflection points of 8 out of the 15 morphometric characters studied. Additionally, temperature significantly affected the allometry coefficients, with a decreasing growth intensity as the temperature difference decreased. The rate of TL growth increased under elevated temperature conditions throughout the entire ontogenetic period, except during the early larval period (feeding onset to metamorphosis onset), at which time D. labrax presented equal growth rates at 15°C and 20°C. The results are discussed with respect to the ontogeny of the functional morphology and the meanings of temperature-induced ontogenetic plasticity for the survival of fish larvae.     

Otolith record of age, growth, and ontogeny in larval and pelagic juvenile Stephanolepishispidus (Pisces: Monacanthidae)

Juveniles of the planehead filefish Stephano-lepishispidus (Pisces: Monacanthidae) (Linnaeus, 1766) are a major component of the Sargassum spp. community, yet little is known of their ecology. In this study, the otolith record of age, growth, and ontogeny in S. hispidus was examined. Juveniles caught off Beaufort Inlet, North Carolina (USA) on 30 June 1996 were marked with alizarin complexone and reared in a flow-through, outdoor tank for up to 19 days. Examination of marked otoliths at several time intervals showed that increment formation was not significantly different than one increment per day, and thus, increment number was used to estimate age. Depth-distribution, morphology, and meristics of larvae and juveniles collected (1990–1992) between Cape Romain, South Carolina, and Cape Hatteras, North Carolina, were examined to identify the timing of the larval to juvenile transition. All indicators suggested the transition occurred between 17 and 20 days. Mean otolith increment widths exhibited a marked change at about 20 days, coinciding with the timing of the larval to juvenile transition and a change in the depth distribution from bottom to surface waters. Increment width of individual juveniles, however, did not exhibit the same pattern; only 40% conformed to the pattern identified for all fish. Thus, the record of the larval to juvenile transition is clear at the population level, but unresolved at the individual level. Received: 1 November 1999 / Accepted: 18 December 2000     

Influence of development and rearing temperature on the distribution,ultrastructure and myosin sub-unit composition of myotomal muscle-fibre types in the plaice Pleuronectes platessa

Eggs of the plaice Pleuronectes platessa L. were incubated at temperatures of 5, 8, 10, 12 and 15°C in March 1990, 1991 and 1992. The myotomes of yolk-sac larvae contain a single superficial layer of small-diameter muscle fibres which stain intensely for succinic dehydrogenase activity, surrounding 390 to 500 weakly staining inner-muscle fibres of larger diameter. Larvae reared at 15°C only survived for a few days and had significantly more inner-muscle fibres of larger average cross-sectional area than those hatching at 5 to 10°C. Myofibrils occupied 61% of the volume of inner-muscle fibres in 15°C larvae compared with 35 and 36% in larvae hatching at 5 and 10°C, respectively (P(0.01). Following metamorphosis, which occurs between 7 and 10 wk, the myotomes retain the single layer of superficial-muscle fibres characteristic of larvae. A thickening of the superficial-muscle layer is first evident in 4 to 5 mo-old laboratory-reared fish of 20 mm total length (TL) and in 0-group fish caught in June and July. On the basis of the histochemical staining reactions for myofibrillar ATPase and succinic dehydrogenase activities, the myotomes of 1-group (104 mm TL) and adult (280 mm TL) plaice were found to contain a minimum of six distinct muscle-fibre types. Two-dimensional gel electrophoresis and peptide mapping were used to investigate changes in myosin subunit composition during development. Myosin from the inner muscle of larvae contains two isoforms of the phosphorylatable light-chain 2 (LC2_L1 and LC2_L2). Following metamorphosis and during the first year, inner-muscle fibres co-express LC2 isoforms characteristic of the superficial fast-muscle fibres of adult plaice (LC2_F1 and LC2_F2) in addition to the larval isoforms. Fast-muscle fibres isolated from deep layers of the myotomes in adult plaice only contain LC2_F2. In contrast, myosin from larval muscle and adult fast muscle contain apparently identical alkali light chains (LC1 and LC3). Peptide maps of myosin heavy chains (MHCs) from 6 wk-old larvae and 10 wk-old fish that had completed metamorphosis are similar, but distinct from those of 1-group plaice. Further changes in white-muscle MHC composition are evident between 1-group fish of 104 mm TL and adults of 280 mm TL.     

Microstructure of settlement-marks in the otoliths of tropical reef fishes

The morphology and ultrastructure of the otolith settlement-mark was examined in 44 tropical reef-fish species spanning nine families. A classification scheme based on similar otolith characteristics is presented. Three major categories are identified based on changes in increment width and optical qualities of the settlement-mark. Of the 44 species examined, 39 possessed “abrupt” settlement-marks (Type I) characterised by a rapid decrease in increment width (up to 50% reduction) over settlement. Type I settlement-marks were found in all nine families examined. The 39 species spanned the whole range of possible larval durations (Pomacentrus moluccensis, 15 d ± 0 SE; Naso hexacanthus, 91.2 d ± 2.97 SE). Four of the 44 species possessed “zonal” settlement-marks (Type II), featuring a band of increments that are wider than pre-settlement increments. Species in this category are the labrids Corisaygula, Thalassoma bifasciatum, T. lunare and an unidentified acanthurid (Acanthurus sp. 2). One species of acanthurid (N. brevirostris) possessed a “gradual” settlement-mark (Type III), manifest as a gradual decrease in increment width during the settlement period. A possible fourth type was identified from the literature. Gnatholepis thompsoni and Coryphopterus glaucofraenum possessed a settlement-mark with increment widths that increased post-settlement. Available data suggest a poor relationship between the structure of the settlement-mark and the magnitude of metamorphosis (previously reported as internal and external morphological change). Evidence suggests that the increment profile over early development and the increment transitions associated with the settlement event are taxon-specific and may enable late-larval stage fishes to be identified to species level. Received: 21 May 1997 / Accepted: 3 February 1999     

Early life history and recruitment of the tropical eel Anguilla bicolor pacifica, as revealed by otolith microstructure and microchemistry

Otolith microstructure and microchemistry of the tropical eel Anguilla bicolor pacifica Schmidt were examined in glass eels collected at the mouth of the Dumoga River, North Sulawesi Island, Indonesia. Ages of the glass eels examined (age at recruitment) ranged from 124 to 202 d (167 ± 19.3 d; mean ± SD), hatching being estimated as having occurred between November 1995 and March 1996. Otolith increment widths markedly increased from age 101 to 172 d (135 ± 18.2 d; mean ± SD), coincident with a drastic decrease in otolith Sr:Ca ratios, suggesting that metamorphosis began during that period. The duration of metamorphosis was estimated as 20 to 40 d, on the basis of otolith microstructural characteristics. The fluctuation patterns in otolith increment widths and Sr:Ca ratios were similar to those of the temperate Japanese eel A. japonica. Received: 20 May 1998 / Accepted: 7 October 1998     

Ontogeny of osmoregulation in the European sea bass Dicentrarchus labrax L.

 Under natural conditions, sea bass eggs hatch in the open sea and young-of-the-year sea bass are found close to estuaries and enter brackish and even freshwater lagoons in the Mediterranean and the eastern Atlantic. The ontogeny of osmoregulation in the European sea bass, Dicentrarchus labrax (Linnaeus, 1758), was studied in different developmental stages, from hatching to large juveniles, exposed to a range of salinities at 17 °C. The experiments were carried out from March to August 1998 in southern France. The type of hyper-hypo-osmotic regulation did not change during development. All stages hyper-regulated at low salinity (under 10–11‰) and hypo-regulated at higher salinities. The acquisition of the full ability to hypo-regulate occurred in four steps. Osmoregulatory capacity was size- and age-dependent and reached its maximum for fish 17–26 mm long, 63–86 days after hatching. The iso-osmotic salinity was 10.2–11.6‰. Our results suggest that early development of osmoregulatory ability, and thus of salinity tolerance in sea bass, may provide an advantageous flexibility for the timing of migration towards low-salinity habitats. Received: 22 May 2000 / Accepted: 18 December 2000     

Halogenated metabolites in two marine polychaetes and their planktotrophic and lecithotrophic larvae

This study investigated the occurrence and ontogenetic changes of halogenated secondary metabolites in planktotrophic and lecithotrophic larvae and adults of two common, infaunal polychaetes, Streblospio benedicti (Spionidae) and Capitella sp. I (Capitellidae), with different life-history traits. S. benedicti contains at least 11 chlorinated and brominated hydrocarbons (alkyl halides) while Capitella sp. I contains 3 brominated aromatic compounds. These halogenated metabolites are potential defense compounds benefiting both larvae and adults. We hypothesized that: (1) planktotrophic larvae contain halogenated metabolites because they are not protected by adult defenses, (2) quantitative and qualitative variation of planktotrophic larval halogenated metabolites parallels that of adults, and (3) brooded lecithotrophic larvae initiate the production of halogenated metabolites only after metamorphosis. To address these hypotheses, volatile halogenated compounds from polychaete extracts were separated using capillary gas chromatography and identified and quantified using mass spectrometry with selected ion monitoring. All four life stages (pre- and post-release larvae, new recruits, adults) of both S. benedicti and Capitella sp. I contained the halogenated metabolites previously identified from adults. This is the first report of halocompounds identified and quantified in polychaete larvae. Allocation of potential defense compounds to offspring varied as a function of species, feeding type and developmental stage. Pre-release larvae of S. benedicti with planktotrophic development contained the lowest concentration of total halogenated metabolites (1.75 ± 0.65 ng mm⁻³), post-release and new recruits contained intermediate concentrations (8.29 ± 1.72 and 4.73 ± 2.63 ng mm⁻³, respectively), and planktotrophic adults contained significantly greater amounts (28.9 ± 9.7 ng mm⁻³). This pattern of increasing concentrations with increasing stage of development suggests synthesis of metabolites during development. Lecithotrophic S. benedicti post-release larvae contained the greatest concentrations of halometabolites (71.1 ± 10.6 ng mm⁻³) of all S. benedicti life stages and developmental types examined, while the amount was significantly lower in new recruits (34.0 ± 15.4 ng mm⁻³). This pattern is consistent with a previously proposed hypothesis suggesting a strategy of reducing potential autotoxicity during developmental transitions. Pre-release lecithotrophic larvae of Capitella sp. I contained the highest concentration of total halogenated metabolites (1150 ± 681 ng mm⁻³), whereas the adults contained significantly lower total amounts (126 ± 68 ng mm⁻³). All concentrations of these haloaromatics are above those known to deter predation in previously conducted laboratory and field trials. As a means of conferring higher larval survivorship, lecithotrophic females of both species examined may be expending more energy on chemical defenses than their planktotrophic counterparts by supplying their lecithotrophic embryos with more of these compounds, their precursors, or with energy for their synthesis. This strategy appears common among marine lecithotrophic larval forms. Received: 14 July 1999 / Accepted: 20 January 2000