Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae)

The plankton larval duration for 100 species of Pacific and Atlantic damselfishes was estimated from daily growth increments on the otolith of juvenile fish collected at various localities between July 1987 and September 1988. For newly-settled fishes, larval duration was determined by counting the entire number of increments present on the otolith, while for older juveniles estimates were made by counting the number of increments between the center of the otolith and a mark corresponding to settlement. We document the development of otolith formation during the period when eggs are incubated on the reef and show that daily increments are only accreted after larvae hatch and enter the planktonic phase. The planktonic larval duration for damselfish is shorter and less variable, both between and within species, compared to other groups of reef fishes such as wrasses and surgeonfishes. Larval duration ranged from 12 to 39 d. Average duration between species ranged from 13.1 to 35.2 d. The time spent in the plankton was not significantly correlated with geographic distribution when evaluated among species, however, genera with confined regional distribution have a shorter mean larval life than do widely distributed genera. Size at settlement was positively correlated with time spent in the plankton among species, but a significant correlation between these variables was only evident within one of ten species. The low variance in planktonic larval duration within species indicates that most damselfish are unable to delay metamorphosis following competency. This inability to postpone settlement limits the potential for dispersal, especially when dispersal time between suitable habitats is greater than about 30 d.     

Settlement patterns and post-settlement survival in two Mediterranean littoral fishes: influences of early-life traits and environmental variables

We examined the relationships between daily pattern of settlement and environmental parameters during two consecutive years in two littoral fishes, Lipophrys trigloides (Blenniidae) and Chromis chromis (Pomacentridae), in the NW Mediterranean Sea. We also used individual early-life traits (pelagic larval duration, size at hatching and size at settlement) calculated from otoliths, to study the proximate causes of settlement variability and size-selective mortality after settlement. Several early-life characteristics of L. trigloides (planktonic larval duration and size at hatching), and environmental variables averaged during the whole planktonic period (e.g. water temperature, wave height, solar radiation) were related with the magnitude of settlement. In contrast, C. chromis showed no significant relationships between early-life traits and the magnitude of settlement, and a weak relationship between settlement magnitude and environmental variables. Furthermore, juvenile survivors showed larger size at hatching than settlers, indicating that size at hatching affected the juvenile survival of the two species. These results suggest that survival was linked largely to conditions at hatching for both species.     

Growth,dispersal, and identification of planktonic labrid and pomacentrid reef-fish larvae in the eastern Pacific Ocean

Planktonic larvae of six genera of labrid and pomacentrid reef fishes were captured in march 1985 in the eastern Pacific Ocean several hundred kilometers from the nearest reefs. The larvae were identified to genus by fin-ray counts as well as by comparison of their larval otolith morphology with that of known species. The larval otolith morphologies of known species were derived from measurements of the larval otolith embedded within the otoliths of settled juveniles (as delineated by the daily otolith-increment marks corresponding to the late larval period). The body morphology and melanophore patterns of the eastern Pacific labird and pomacentrid larvae closely matched those of congeneric larvae described from other oceans. Growth rates of larvae less than about 70 d old were similar between taxa (from 0.13 to 0.19 mm d^-1). After about 70 d in the plankton, labrid larvae grew much more slowly (0.06 mm d^-1 in Xyrichtys sp.). Labrid larvae had long larval durations (up to 131 d in Xyrichtys sp.), while the larval lives of the pomacentrids appeared to be shorter and much less variable. Larvae of many different ages occurred within the same water mass, and young cohorts of larvae appeared continuously over the sampling period. Some larvae were as young as 21 d, indicating that reef-fish larvae are capable of rapid long-distance dispersal (at least 18 km d^-1).     

Daily otolith increments and recruitment in two coral-reef wrasses,Thalassoma bifasciatum and Halichoeres bivittatus

Increments on the otoliths of two common coral reef fishes, the bluehead wrasse Thalassoma bifasciatum and the slippery dick Halichoeres bivittatus, were demonstrated by mark-recapture experiments to be daily. Otoliths were marked in two ways; by depriving fish of light, food, and temperature cycles and also by supplemental feeding in the field. Both experiments were performed in late 1980 in the San Blas Islands of Panamá. A mark corresponding to settlement of the planktonic larva onto the reef was found on the otoliths of the bluehead wrasse. This settlement mark was used to calculate the dates of settlement of a collection of juveniles of this species taken from a patch reef in the San Blas Islands of Panamá in 1981. Settlement occurred in short and irregular bursts. The number of daily increments before the settlement mark indicates a planktonic larval life of 40 to 72 d.     

Latitudinal variation in abundance of herbivorous fishes: a comparison of temperate and tropical reefs

The aim of the study was to provide comparable estimates of abundance of herbivorous reef fishes at temperate and tropical localities using a standardized methodology. Faunas of herbivorous fish were sampled on the rocky reefs of temperate northern New Zealand and on the coral reefs of the northern Great Barrier Reef (GBR), Australia, and the San Blas Archipelago in the Caribbean. A pilot study established the most appropriate habitat setting and the scale and magnitude of replication for the sampling program in temperate waters. Herbivorous fishes, including members of families endemic to the southern hemisphere (Odacidae and Aplodactylidae), were most abundant in turbulent, shallow water (0 to 6 m) and had patchy distributions within this habitat. A hierarchical sampling program using 10-min transect counts within the 0 to 6 m depth stratum examined abundance patterns at a range of spatial scales including mainland and island coasts, localities separated by up to 100 km and sites separated by up to 10 km. This program identified a characteristic fauna of seven species of herbivorous fishes with mean total abundances ranging from 23 to 30 individuals per 10-min transect. Species composition of the fauna varied between islands and coasts. A similar methodology was used to sample the major families of herbivorous fish in a number of sites in each of the tropical regions. These sampling programs revealed a fauna dominated by acanthurids and scarids in both the GBR and Caribbean localities. Estimates of abundance from these regions were similar, with a mean of 108 individuals recorded on the GBR and 129 per 10-min transect in the Caribbean. Species richness varied between each region, with 44 taxa recorded from the GBR and 11 from the Caribbean. Abundances of temperate water herbivores in New Zealand were found to be 75 to 80% lower than those recorded from shallow water habitats sampled on coral reefs. This was not related to species richness, since both New Zealand and the Caribbean locality had patterns of low richness. We suggest that the differences in abundance found by our study between temperate and tropical regions are not restricted to herbivorous fishes, but are representative of general latitudinal trends in reef fish faunas. Received: 4 November 1996 / Accepted: 15 December 1996     

Exploring the potential of otolith microchemistry to enhance diet analysis in pinnipeds

Increasing the scope and accuracy of information about pinniped diets obtainable from non-invasive techniques is increasingly important, particularly in cases where pinniped species are threatened or endangered. This study is the first to explore the potential for using elemental analysis of the otoliths found in scat to enhance the information available for diet analyses. We investigated the effects of pinniped digestion on otolith microchemistry using laser ablation inductively coupled plasma mass spectrometry (LA-ICPMS). We compared the elements contained in the edges (adult stage) and cores (larval/juvenile stage) of otoliths from Pacific herring (Clupea pallasii), Atka mackerel (Pleurogrammus monopterygius), and Walleye pollock (Theragra chalcogramma) recovered from the scat of captive Steller sea lions (Eumetopias jubatus) to elements in a sample of pristine (undigested) otoliths. We found that digestion had a significant effect on four of the six sampled combinations of species and otolith region (herring edges and cores, mackerel edges, and pollock cores), and that Rb most frequently showed significant differences in concentration after digestion. We could significantly discriminate among species of both pristine and digested otoliths using either otolith edges or cores with the elements Ba, Rb, Sr, Y, and Mg. When compared to previously identified digested otoliths, unknown samples of the three species of digested otoliths could be discriminated with 55–100% accuracy depending on species and otolith region. When compared to a library of previously identified pristine and digested otoliths, unknown samples of digested otoliths could be discriminated to species with 65–88% accuracy. When the group of unknown digested otoliths was compared to known pristine otoliths, discrimination ranged from 45 to 65%. These results indicate that elemental analysis could be used to supplement visual identification of otoliths from scat. However, further research is required to determine whether elemental analysis of digested otoliths could be useful for prey fish population studies.     

Homing to scars as a defense against predators in the pulmonate limpet Siphonaria gigas (Gastropoda)

The length of larval life in some coral reef fishes was estimated from the number of growth increments in the otoliths of newly settled fishes. We examined 210 individuals comprising 38 species and 5 unidentified taxa, and belonging to 12 families. During 2 successive austral summers (1976–1977 and 1978–1979), specimens were collected from the lagoon at One Tree Reef, Great Barrier Reef, Australia. By assuming that growth increments in otoliths are laid down on a regular daily cycle commencing near the time of hatching, we calculated typical ages ranging from 3 to 6 wk with a minimum of just over 2 wk and a maximum of 12 wk. The otoliths also contain distinctive microstructural features which can serve as approximate temporal markers for the change from the postlarval to juvenile life stage.     

Regional differences in duration of the planktonic larval stage of reef fishes in the eastern Pacific Ocean

Regional variation in the duration of the planktonic larval phase of three species of reef fishes, Thalassoma lucasanum (Labridae), Stegastes flavilatus, and Microspathodon dorsalis (Pomacentridae) was investigated between 1982 and 1991 at several sites in the tropical eastern Pacific over a distance of 3500 km, encompassing virtually their entire range of distribution. Durations of the larval phase, determined from counts of daily otolith increments, were significantly different (1.3 to 1.6 x) between sites. Populations of all three species had a consistently shorter larval life at the most northern site, Cabo San Lucas (Mexico) compared to Panamá and the offshore islands of Galápagos and Cocos. Analyses of otolith increment width over the precompetent period revealed that this disparity in larval duration primarily reflected differences in larval growth rates: faster growing fish spent less time in the plankton. In T. lucasanum, some of the variation in larval duration between Panamá and offshore sites (Galápagos Islands and Cocos Island) may be accounted for by a higher frequency of individuals delaying metamorphosis at the offshore sites. These data indicate that conditions in the planktonic environment are not homogeneous throughout the tropical eastern Pacific and may have a profound effect on aspects of the larval ecology of reef fishes in this region.     

Climatic effects on the growth of a temperate reef fish from the Southern Hemisphere: a biochronological approach

Increments in the hard parts of marine organisms (otoliths, skeletons, shells) can provide long-term chronologies of growth analogous to tree rings. For the first time in the Southern Hemisphere, we use a dendrochronological (tree-ring analysis) approach to develop a multidecadal chronology of growth for a temperate reef fish, Girella tricuspidata, from the coast of northern New Zealand. Growth patterns in the otoliths of this species were strongly synchronous among individual fish over a period spanning 27 years (1980–2006). We then compared our otolith chronology to climatic records and found strong positive correlations of growth with sea surface temperature, and weak negative correlations with the multivariate El Nino Southern Oscillation (ENSO) index. Strongest correlations were found between summer sea surface temperature and otolith growth. This relationship was consistent across all years and explained 44 % of the variation (y = −2.0 + 0.1785 × temperature, r ² = 0.4367, P = 0.0002) in the G. tricuspidata growth chronology. Our study illustrates how otolith chronologies provide remarkable records of annual growth patterns over decadal time scales that will be useful for forecasting the likely effects of climate change on marine ecosystems.     

Otolith size-at-hatch reveals embryonic oxygen consumption in the zebrafish, <Emphasis Type="Italic">Danio rerio</Emphasis>

Otoliths have frequently been used to reconstruct growth histories in larval, juvenile and adult fish. However, there is growing evidence that otolith growth is directly determined by metabolic intensity and, consequently, only indirectly related to somatic growth. By performing measurements of oxygen consumption rate and other early life-history traits on individual eggs of zebrafish (Danio rerio), we found that oxygen consumption explained residual variance in otolith size that is not accounted for by egg size. Total oxygen consumption during the embryonic stage explained 34% of the variance in sagitta size at hatch, whereas larval size at hatch (as a proxy for growth during the embryonic period) was not significantly correlated with sagitta size. This strongly suggests that otolith growth is directly related to metabolic rate, and yields a mechanism that may explain recent observations of a link between otolith size at hatch and viability in larval fish.     

Critical swimming speeds of late-stage coral reef fish larvae: variation within species,among species and between locations

The swimming abilities of larval fishes are important for their survival, potentially affecting their ability to avoid predators, obtain food and control dispersal patterns. Near settlement swimming abilities may also influence spatial and temporal patterns of recruitment. We examined Critical speed (U-crit) swimming ability in late stage larvae of 89 species of coral reef fishes from the Great Barrier Reef and the Caribbean. Coefficients of variation in U-crit calculated at the individual level were high (28.4%), and this was not explained by differences in size or condition factor of these same larvae. Among species U-crit ranged from 5.5 cm s⁻¹ to 100.8 cm s⁻¹ (mean=37.3 cm s⁻¹), with 95% of species able to swim faster than the average current speed around Lizard Island, suggesting that most species should be capable of influencing their spatial and temporal patterns of settlement. Inter-specific differences in swimming ability (at both the family and species levels) were significantly correlated with size and larval morphology. Correlations were found between swimming performance and propulsive area, fineness ratio and aspect ratio, and these morphological parameters may prove useful for predicting swimming ability in other taxa. Overall, the swimming speeds of larvae from the same families at the two locations were relatively similar, although the Lutjanidae and Acanthuridae from the Caribbean were significantly slower than those from the great barrier reef. Differences in swimming speed and body form among late stage larvae suggests that they will respond differently to factors influencing survival and transport during their pelagic phase, as well as habitat use following settlement.     

In situ experiment of ontogenetic variability in the otolith chemistry of Stegastes partitus

Otolith chemistry can be used to assess pelagic larval fish connectivity by comparing spatially variable otolith edge chemistry (corresponding to the site of collection) to otolith core chemistry (corresponding to the site of hatching). However, because the otolith’s edge and core represent different life stages, the deposition of elements may differ, thus complicating direct comparisons of edge and core chemistry to investigate connectivity. Here we present data from a field experiment in which otoliths from embryos (3 days post-fertilization) and juveniles of Stegastes partitus were collected at the same site and time, and chemically analyzed to assess whether elemental concentrations of otoliths vary ontogenetically. Separate multivariate analyses, each investigating the spatial/temporal variability in the chemistry of either embryo otoliths or the edges of juvenile otoliths, revealed significant differences, suggesting an environmental influence to the chemical signals of otoliths. A nested multivariate analysis assessing whether otolith chemistry varied with life history stage (i.e., ontogenetic variability) indicated that elemental concentrations of embryo otoliths were significantly greater than that of juvenile otolith edges. Specifically, embryo elemental concentrations of Mn, Zn, Sn, Ba, Ce, and Pb were between 2 and 163 times greater than those of the corresponding juvenile otoliths, and thus the environment was not the primary determinant of embryo otolith chemistry. Consequently, caution is warranted when interpreting environmental patterns of otolith cores, particularly when using them as a proxy for natal signatures.     

Effects of selective settlement and of aggression by residents on distribution of young recruits of two tropical damselfishes

The reef fishes that settled on an array of experimental corals at Lizard Island, Queensland, Australia, were counted during a pulse of recruitment in December 1986. NeitherPomacentrus sp. norP. amboinensis showed any evidence that harassment by residentDascyllus aruanus caused a decrease in persistence during the first day after settlement. LarvalPomacentrus sp. settled selectively on corals without residentD. aruanus. The results forP. amboinensis were ambiguous. Settlers of both species positioned themselves closer to the sand on corals with residentD. aruanus than on unoccupied corals. This could reduce access to planktonic food and increase the risk of predation. Adult aggression may be less important and active selection of settlement sites by larvae may be more important to the distribution of recruits than is suggested by the literature. The presence or absence of particular species should be included among the cues that larval reef fishes use to choose settlement sites.     

Elevated manganese concentrations at the cores of clupeid otoliths: possible environmental,physiological, or structural origins

Laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS) was used to investigate the distribution of manganese (Mn) in Atlantic herring (Clupea harengus L.) otoliths. An unusual pattern of Mn concentration, which has not previously been reported in fish otoliths, was observed. Time-resolved plots (time roughly equivalent to ablation depth) of the Mn signal recorded in the ICP-MS detector during ablation of otoliths contained a distinct peak, suggesting that elevated Mn concentrations were specific to certain regions. Ablation inside and outside of the otolith core revealed that the region of high Mn concentration coincided with the ablation of the core. It was initially suggested that Mn from the gravel substrate on which herring eggs are incubated is incorporated into the otolith core, producing elevated Mn concentrations in this region. It was hypothesised that otoliths from fish whose eggs are not incubated in gravel will not contain high Mn concentrations at the core. To test this, otoliths of sprat (Sprattus sprattus), which have a pelagic embryological stage, and from reared herring, which were incubated on glass plates, were analysed using LA-ICP-MS. Peaks in the Mn signal were observed during the ablation of cores from some sprat and reared herring otoliths, indicating that incubation on a gravel substrate is not necessary for the accumulation of Mn at the otolith core and that otoliths from both pelagic and demersally spawning fish species can contain elevated Mn concentrations. The results suggest that Mn concentrations at the otolith core do not reflect the external environment of the developing embryo. Two other hypotheses are considered. A dramatic increase in Mn concentrations in the egg, associated with embryological development, may produce elevated Mn levels at the otolith core. Alternatively, the high levels of Mn observed at the otolith core may reflect spatial variation in the crystal structure of the calcium carbonate portion of the otolith. The possible influence of physiological or structural factors on the chemical composition of the otolith core may confound the environmental signal in this region of the otolith. This has important implications for the interpretation of otolith chemistry data.Communicated by J.P. Thorpe, Port Erin     

Confirmation of longevity in Sebastes diploproa (Pisces: Scorpaenidae) from 210Pb/226Ra measurements in otoliths

Fish ages are often estimated by assuming an annual frequency of the band-like, growth-zones recorded in the largest of their otoliths, the sagittae. The total number of growth-zones are normally determined either by counting external growth-zones (whole otolith technique) or by examining otolith cross-sections (otolith section technique). The two techniques do not always yield the same age, however, particularly in older specimens of certain fishes. To resolve this problem, otoliths of the splitnose rockfish Sebastes diploproa were examined morphologically and were assayed for their natural radionuclide concentrations. Four age groups of otoliths were identified based on growth-zone counting; in the first three, whole otolith and otolith section age estimates agreed, while in the fourth, the otolith section age substantially exceeded the whole otolith age. Radiometric analysis demonstrated that all otoliths were deficient in ²¹⁰Pb activity relative to ²²⁶Ra activity with the deficiency decreasing with increasing number of growth-zones. The magnitude of the ²¹⁰Pb/²²⁶Ra radioactive disequilibrium in each otolith group, when compared to the number of growth-zones and the otolith weight histories derived with the two techniques, identifies the growth-zones revealed by otolith sections as annual features. Thus when otolith section age exceeds whole otolith age (usually occurring after 20 to 25 yr of age for this species), the otolith section technique is the correct method of age determination. Estimates of longevity in the genus Sebastes near 80 yr are therefore confirmed.     

Sclerochronological studies reveal that patterns of otolith growth of adults of two co-occurring species of Platycephalidae are synchronised by water temperature variations

Chronologies are developed from the otolith growth-increment widths of adult rock flathead (Platycephalus laevigatus) and longhead flathead (Leviprora inops) collected from three inter-connected embayments in temperate south-western Australia. Marginal increment trends on otoliths, in combination with the dendrochronological technique of crossdating, provide strong evidence that an opaque zone is formed annually in the otoliths of both species. Increment widths between opaque zones in the otoliths of individuals of P. laevigatus (maximum age 20 years) and L. inops (maximum age 21 years) were synchronised within and between species. The correlation between the two master otolith chronologies, i.e. mean standardised increment width for each year, was highly significant (P < 0.01). The master otolith chronology for each species was positively correlated with mean sea surface temperatures from 1 July to 30 June, which encompassed the austral summer when growth is typically greatest, and with mean monthly sea surface temperature between September and March. Leeuwin Current strength, which influences the biotic characteristics of some marine species on the south coast, has little influence on the pattern of otolith growth, presumably because the inter-connected embayments from which fish were obtained were partially enclosed. On the basis of the above data and the geographical distributions of P. laevigatus and L. inops, it is proposed that, while both species would grow faster with increasing temperature, the temperatures on the south coast are closer to the optimum for the former species.     

Population genetic structure of the neon damselfish (<Emphasis Type="Italic">Pomacentrus coelestis</Emphasis>) in the northwestern Pacific Ocean

The population genetic structure of the neon damselfish (Pomacentrus coelestis) in the northwestern Pacific Ocean was revealed by the hypervariable control region of the mitochondrial gene (343 bp). In total, 170 individuals were sampled from 8 localities distributed between Taiwan and Japan, and 71 haplotypes were obtained through sequence alignment. High haplotype diversity (h = 0.956 ± 0.008) with low nucleotide diversity (π = 0.010 ± 0.006) was observed, and the results of the mismatch distribution test suggested that a historical population expansion after a period of population bottleneck might have occurred among P. coelestis populations. Based on the results of the UPGMA tree and AMOVA (Φct = 0.193, P < 0.05) analyses, fish populations from eight localities could be divided into two groups: one includes populations from localities around mainland Japan, and the other includes those from Okinawa and southern Taiwan. A genetic break was found between populations from mainland Japan and Okinawa, and this break was congruent with the pattern of phenotypic variations documented in previous studies. This evidence supports the latitudinal variation of reproductive traits among P. coelestis populations likely being genetically based. It is suggested that the changes in sea level and sea surface temperatures during past glaciations might have resulted in population bottlenecks in P. coelestis and the modern populations in the northern West Pacific are likely the results of recolonization after such events. The Kuroshio Current acts not only as a vehicle for larval transport along its pathway (between populations in southern Taiwan and Okinawa) but also as a barrier for larval dispersal across the Kuroshio axis (between populations in mainland Japan and Okinawa). Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Early life history of the American conger eel (<Emphasis Type="Italic">Conger oceanicus</Emphasis>) as revealed by otolith microstructure and microchemistry of metamorphosing leptocephali

The early life history of the American conger eel, Conger oceanicus, was studied using otolith microstructure and chemical composition in metamorphosing leptocephali collected from New Jersey estuarine waters. The age of leptocephali was estimated by counting daily growth increments. Age of early metamorphosing leptocephali at recruitment to the estuary ranged from 155 to 183 days, indicating that migration of conger eel leptocephali from their oceanic spawning ground to the estuary requires 5–6 months. Back-calculated hatching dates suggest that the spawning season lasted 3 months, from late October to mid-December. However, in the late metamorphic leptocephali, the presence of an unclear peripheral zone in the otolith prevents the accurate estimation of the larval stage duration. The calcium content was almost constant throughout the otoliths. Both strontium and Sr:Ca ratios increased with age, but dramatically decreased at age 70–120 days. The otolith increment width also showed a marked increase at the same ages, indicating the onset of metamorphosis. A negative correlation between age at metamorphosis and otolith growth rate indicates that faster growing leptocephali arrive at the estuary earlier than slower growing ones. A close relationship was also found between age at recruitment and age at metamorphosis, suggesting that individuals that metamorphosed earlier were recruited to the estuary at a younger age. This larval migration pattern appears to be similar among anguilliform fishes.Communicated by S.A. Poulet, Roscoff