Caging experiment to examine mortality during metamorphosis of coral reef fish larvae

All previous attempts to estimate early postsettlement mortality of coral reef fishes using either caging experiments or disappearance of new recruits have examined fish that had already settled, and therefore did not include the metamorphosis process. Crest nets capture unharmed transparent larvae during their migration from the open ocean to lagoon reefs before metamorphosis. We released these presettlement larvae at night into cages surrounding patch reefs and measured larval survivorship after two nights. This caging experiment involved cages enclosing the natural resident fish fauna, including predators, and others cleared of fish before releasing the larvae. The analyses of variance showed that (1) there was no difference in survivorship between the seven trials, (2) there was a significant difference between cleared and uncleared cages, and (3) there were significant differences between larval species tested. For the seven species that had a significant difference in survivorship between cleared and uncleared cages, average mortality of the larvae was 14% (range 0-26%) in cleared cages and 67% (range 29-76%) in cages with predators. The difference in mortality between species was related to the size of the larvae, as larger species exhibited reduced mortality compared to smaller species. Mortality was related to the abundance of resident fish that could act as predators or competitors. Predation can have a significant impact on the survival of metamorphosing fish larvae on coral reefs.     

Temperature influences swimming speed, growth and larval duration in coral reef fish larvae

The effects of temperature on growth, pelagic larval duration (PLD) and maximum swimming speed were compared in the tropical fish marine species Amphiprion melanopus, to determine how temperature change affects these three factors critical to survival in larvae. The effects of rearing temperature (25 and 28 °C) on the length of the larval period and growth were examined in conjunction with the effects of swimming temperature (reared at 25 °C, swum at 25 and 28 °C, reared at 28 °C, swum at 25 and 28 °C) on critical swimming speed (U-crit). Larvae reared at 25 °C had a 25% longer pelagic larval duration (PLD) than larvae reared at 28 °C, 12.3 (±0.3) days compared with 9 (±0.6) days at 25 °C. To offset this effect of reduced developmental rate, growth and U-crit were measured in larvae reared at 28 and 25 °C at the same absolute age (7 days after hatching (dah)) and same developmental age (7 dah at 28 °C cf. 11 dah at 25 °C), corresponding to the day before metamorphosis. Larvae reared at 25 °C were smaller than larvae reared at 28 °C at the same absolute age (7 dah at 25 °C cf. 7 dah at 28 °C), yet larger at similar developmental age (11 dah at 25 °C cf. 7 dah at 28 °C) when weight and standard length were compared. This stage-specific size increase did not result in better performance in larvae at the same developmental age, as there was no difference in U-crit in premetamorphic larvae reared at either temperature (7 dah at 28 °C c.f 11 dah at 25 °C). However, U-crit was considerably slower in 7-day-old larvae reared at 25 °C than larvae of the same absolute age (7 dah) reared at 28 °C. Swimming temperature controls demonstrated that a change in temperature immediately prior to swimming tests did not effect swimming performance for larvae reared at either temperature.A decreased in rearing temperature resulted in longer larval durations, reduced growth rates and slower swimming development in larvae. However, the magnitude of the response of each of these traits varied considerably. As such, larvae reared at the lower temperature were a larger size at metamorphosis but had poorer relative swimming capabilities. This study highlights the importance of measuring a range of ecologically relevant traits in developing larvae to properly characterise their relative condition and performance in response to environmental change.     

Larval mortality and the composition of coral reef fish communities

Research over the past decade shows that fish populations on coral reefs can vary enormously, both spatially and temporally. Nonetheless, predictable patterns in structure are present at both small and regional scales. These have usually been interpreted as resulting from processes acting after settlement of fishes from the plankton. However, current research now suggests that planktonic processes could also result in deterministic patterns of community structure.     

Parental condition affects early life-history of a coral reef fish

Parents can exert a range of non-genetic effects on the growth and survival of their offspring. In particular, parents may modify the size or condition of their offspring depending on the amount of energy they have available for reproduction. In this study, the body condition of breeding pairs of the coral reef fish Acanthochromis polyacanthus was experimentally manipulated to test the effects of parental condition on reproductive output and offspring life history traits. Parents in good condition commenced breeding earlier, had higher reproductive output, and their eggs exhibited increased survival during embryogenesis, compared to parents in poorer condition. Increased reproductive output was attained through more reproductive bouts over the breeding season that contained both a greater number and an increased size of eggs. The offspring from parents in good condition were larger at hatching, with larger yolk reserves and increased survival on endogenous reserves. Larger size is expected to provide benefits to offspring through reduced susceptibility to size-selective mortality. The range of offspring characteristics modified by parental condition could result in a greater proportion of offspring from good condition parents recruiting to the population.     

Comparative organ differentiation during early life stages of marine fish

The basic developmental mechanisms of teleosts are similar, but there are differences with respect to the timing of developmental events. These events are controlled by genetic and environmental factors. Direct comparisons of organogenesis are complicated due to large variations in egg sizes and incubation temperatures between species. But in general, cultivated small marine pelagic fish larvae originating from rather small eggs (like gadoids, flatfishes, sparids) hatch with a relatively large yolk sac, a larval finfold and subdermal space and under-developed organs. Developmental status at hatch differs between species and the duration of the yolk sac period varies. Main organs and organ systems become functional by first feeding and differentiate during the larval stage and metamorphosis. Species developing directly via large yolk-rich eggs and a long incubation period have a juvenile like morphology and organ functionality at first feeding, sometimes immediately after hatch (like wolffishes). Histomorphological and cell- or organ functional studies of developing embryos and larvae of cultivated species constitute basic information for understanding species-specific events, of utmost importance for improving production protocols. Information is still lacking on early functionality of endocrine and immunocompetent tissues and organs, areas that deserve future focus.     

Genetic diversity loss associated to high mortality and environmental stress during the recruitment stage of a coral reef fish

We investigated the short-term impact of environmental-induced stress on survival and neutral genetic diversity of recently settled juveniles of a damselfish, Dascyllus aruanus, using spatiotemporal caging experiments in various natural environmental conditions in Moorea (French Polynesia). Juveniles’ mortality was followed at five study sites and overall four experiments, mortality rates ranged from 0 to 45%. Mortality rate and average daily water temperature were positively correlated (P = 0.018). Juveniles’ mortality rate and allelic richness estimated from ten microsatellite loci were negatively correlated (P = 0.046). Together, an overdominance of heterozygotes was observed within hostile environments. These results suggest that an allelic richness loss may be expected as a direct consequence of unfavorable environmental conditions. Thus, a worrisome scenario on demographic and genetic consequences may be expected from habitat degradation in the context of global change and human pressure increases.     

Grazing by a small fish affects the early stages of a post-settlement stony coral

Short-term experiments were used to isolate the detrimental effects of grazer disturbance on young corals, and determine the stage of development at which recruits are no longer susceptible to this disturbance. Artificial substrata containing an algal matrix and coral recruits of different life stages were exposed to grazing by epilithic algal matrix (EAM) feeding combtoothed blennies, Salarias fasciatus. Single polyp recruits were vulnerable to grazer disturbance, while multi-polyp recruits (ca. 6–8 polyps) survived with evidence of minor damage in the form of tissue and polyp loss. The result indicates that blennies, although small and possessing weak dentition, can negatively influence the survival of young coral recruits. The protruding structure of micro-nubbins, representing juvenile corals were not damaged, suggesting that coral achieving that size and form can escape such damage. Communicated by Ecology Editor Prof. Peter Mumby     

Live coral repels a common reef fish ectoparasite

Coral reefs are undergoing rapid changes as living corals give way to dead coral on which other benthic organisms grow. This decline in live coral could influence habitat availability for fish parasites with benthic life stages. Gnathiid isopod larvae live in the substratum and are common blood-feeding parasites of reef fishes. We examined substrate associations and preferences of a common Caribbean gnathiid, Gnathia marleyi. Emergence traps set over predominantly live coral substrata captured significantly fewer gnathiids than traps set over dead coral substrata. In laboratory experiments, gnathiids preferred dead coral and sponge and tended to avoid contact with live coral. When live gnathiids were added to containers with dead or live coral, significantly fewer were recovered from the latter after 24 h. Our data therefore suggest that live coral is not suitable microhabitat for parasitic gnathiid isopods and that a decrease in live coral cover increases available habitat for gnathiids.     

Long-term cleaner fish presence affects growth of a coral reef fish

Cleaning behaviour is considered to be a classical example of mutualism. However, no studies, to our knowledge, have measured the benefits to clients in terms of growth. In the longest experimental study of its kind, over an 8 year period, cleaner fish Labroides dimidiatus were consistently removed from seven patch reefs (61-285 m(2)) and left undisturbed on nine control reefs, and the growth and parasite load of the damselfish Pomacentrus moluccensis determined. After 8 years, growth was reduced and parasitic copepod abundance was higher on fish from removal reefs compared with controls, but only in larger individuals. Behavioural observations revealed that P. moluccensis cleaned by L. dimidiatus were 27 per cent larger than nearby conspecifics. The selective cleaning by L. dimidiatus probably explains why only larger P. moluccensis individuals benefited from cleaning. This is the first demonstration, to our knowledge, that cleaners affect the growth rate of client individuals; a greater size for a given age should result in increased fecundity at a given time. The effect of the removal of so few small fish on the size of another fish species is unprecedented on coral reefs.     

Effects of parasites on larval and juvenile stages of the coral reef fish Pomacentrus moluccensis

The ecological role of parasites in the early life-history stages of coral reef fish is far from clear. Parasitism in larval, recently settled and juvenile stages of a coral reef fish damselfish (Pomacentridae) was therefore investigated by quantifying the ontogenetic change in parasite load and comparing the growth rates of parasitized juvenile fish to those of unparasitized ones. Parasite prevalence in two lunar pulses of Pomacentrus moluccensis was 4 and 0% for larval stage fish, 34 and 56% for recently settled fish and 42 and 49% for juveniles. A significant increase in parasite prevalence with age group was found; the most marked increase occurred immediately after larval fish had settled. Standard length did not model prevalence well; as length is a proxy for age, this indicates that the higher prevalence in recently settled and juvenile fish compared with larvae was not a simple result of parasites accumulating with age. In one of three cohorts, there was some evidence that parasitism affected the growth rate of juveniles, as measured by otolith width. The study suggests that settling on the reef exposes young fish to potentially harmful parasites. This supports the idea that the pelagic phase may have the effect of reducing the exposure of young fish to the debilitating effects of parasites.     

Resolution of the early life history of a reef fish using otolith chemistry

We used the elemental signatures in otoliths of the damselfish Pomacentrus coelestis as a proxy for conditions experienced prior to settlement. Fish from the southern Great Barrier Reef (GBR) differed in their pre-settlement otolith chemistry, indicating that they had occupied different water masses during their pelagic stage, thus suggesting multiple larval sources. Fish from reefs in the northern GBR did not differ greatly in their pre-settlement otolith chemistry, suggesting a single larval source. Using near-natal signatures, we determined that ~67% of these signatures matched the signature established for Lizard Island (LZ), suggesting LZ could be a source reef. However, these results could also be the result of poor separation among reefs caused by reefs sharing water masses. Otolith chemistry also revealed that 50–70% of all fish examined settled on the reef the day they encountered it, while some fish spent up to 4 days near the reef prior to settlement.     

Food availability affects growth in a coral reef fish

Summary Pomacentrus amboinensis is common on small patch reefs within One Tree Lagoon (Great Barrier Reef), where it preferentially settles onto deep reefs. A preliminary experiment, in which juveniles were transplanted to identical reef structures at two sites, within two depth strata, indicated that juvenile growth and survivorship were better in deeper water. The hypothesis that this difference was due to food availability was tested by a supplemental feeding experiment, carried out at another two randomly chosen sites, within the same two depth strata. Fish were fed each day over a one month period, during which no mortality was observed. The growth rates of juveniles were markedly higher on all food-supplemented reefs, when compared to controls. Growth differed between depth strata, but there was no interaction between the food x depth factors, which would have suggested a greater effect of food supplementation in either habitat. Thus, although the difference between depths cannot be attributed to food availability, the results have a more general significance. Food appears to be a limiting resource (in terms of growth) in both the marginal shallow habitat, and the more suitable deeper habitat.     

Patterns of connectivity among populations of a coral reef fish

Knowledge of the patterns and scale of connectivity among populations is essential for the effective management of species, but our understanding is still poor for marine species. We used otolith microchemistry of newly settled bicolor damselfish (Stegastes partitus) in the Mesoamerican Reef System (MRS), Western Caribbean, to investigate patterns of connectivity among populations over 2 years. First, we assessed spatial and temporal variability in trace elemental concentrations from the otolith edge to make a ‘chemical map’ of potential source reef(s) in the region. Significant otolith chemical differences were detected at three spatial scales (within-atoll, between-atolls, and region-wide), such that individuals were classified to locations with moderate (52 % jackknife classification) to high (99 %) accuracy. Most sites at Turneffe Atoll, Belize showed significant temporal variability in otolith concentrations on the scale of 1–2 months. Using a maximum likelihood approach, we estimated the natal source of larvae recruiting to reefs across the MRS by comparing ‘natal’ chemical signatures from the otolith of recruits to the ‘chemical map’ of potential source reef(s). Our results indicated that populations at both Turneffe Atoll and Banco Chinchorro supply a substantial amount of individuals to their own reefs (i.e., self-recruitment) and thus emphasize that marine conservation and management in the MRS region would benefit from localized management efforts as well as international cooperation.     

Experimental confirmation of aggressive mimicry by a coral reef fish

A number of potential mimetic relationships between coral reef fishes have been described, but the underlying mechanisms are poorly understood. Similarities in colour between species have often been attributed to aggressive mimicry (where predators resemble models in order to deceive prey), however this has not been tested. The fang blenny, Plagiotremus rhinorhynchos is a specialized predator that feeds on tissues of other fishes. Some individuals appear to mimic the harmless cleaner wrasse Labroides dimidiatus in order to deceive fish visiting cleaning stations, thereby increasing access to food. In this study, the ecological relationship between the mimic and model was examined at Kimbe Bay (Papua New Guinea) and the hypothesis that colour similarities represent facultative aggressive mimicry was experimentally evaluated. Some juveniles exhibited a striking resemblance to the juvenile colouration of the cleaner wrasse, but only when in close proximity to the wrasse and only when similar in size. As predicted for mimics, P. rhinorhynchos co-occurred with L. dimidiatus, but was rare relative to the model. Among site comparisons showed that the abundance of mimetic type blennies was positively correlated with the abundance of juvenile cleaner wrasses. Approximately 50% of all P. rhinorhynchos were found 1 m from the nearest L. dimidiatus, a distance significantly shorter than expected if they were not associated. A cleaner wrasse removal experiment was carried out to test whether the colour displayed by the blenny and its foraging success were contingent upon the presence of a model. In all cases, removal of the model prompted a rapid colour change to a general non-mimetic colouration in P. rhinorhynchos. Removal of L. dimidiatus also resulted in a ~20% reduction in the average foraging success of the blenny compared to controls, supporting the hypothesis that the blenny is a facultative aggressive mimic of the cleaner wrasse.     

Regulation of local populations of a coral reef fish via joint effects of density- and number-dependent mortality

Density-dependent mortality can regulate local populations - effectively minimizing the likelihood of local extinctions and unchecked population growth. It is considered particularly important for many marine reef organisms with demographically open populations that lack potential regulatory mechanisms tied to local reproduction. While density-dependent mortality has been documented frequently for reef fishes, few studies have explored how the strength of density-dependence varies with density, or how density-dependence may be modified by numerical effects (i.e., number-dependent mortality). Both issues can have profound effects on spatial patterns of abundance and the regulation of local populations. I address these issues through empirical studies in Moorea, French Polynesia, of the six bar wrasse (Thalassoma hardwicke), a reef fish that settles to isolated patch reefs. Per capita mortality rates of newly settled wrasse increased as a function of density and were well approximated by the Beverton-Holt function for both naturally formed and experimentally generated juvenile cohorts. Average instantaneous mortality rates were a decelerating function of initial densities, indicating the per capita strength of density-dependence decreased with density. Results of a factorial manipulation of density and group size indicate that per capita mortality rates were simultaneously density- and number-dependent; fish at higher densities and/or in groups had higher probabilities of disappearing from patch reefs compared with fish that were solitary and/or at lower densities. Mortality rates were ~30% higher for fish at densities of 0.5 fish/m² than at 0.25 fish/m². Similarly, mortality rates increased by ~45% when group size was increased from 1 to 2 individuals per patch, even when density was kept constant. These observations suggest that the number of interacting individuals, independent of patch size (i.e., density-independent effects) can contribute to regulation of local populations. Overall, this work highlights a greater need to consider numerical effects in addition to density effects when exploring sources of population regulation.     

Social transmission of behavioural traditions in a coral reef fish

Traditional behaviours involve the non-genetic transmission of social information across age classes or generations. French grunts (Haemulon flavolineatum) exhibit social traditions of daytime schooling sites and twilight migration routes. Individuals transplanted to new schooling sites and allowed to follow residents at the new sites used the new migration routes and returned to the new sites in the absence of resident fish. Control fish with no opportunity to learn showed no such directionality or return. This is the first demonstration of apparent pre-cultural behaviour in free-living fish. Our observations suggest additional classes of behaviour and taxonomic groups in which pre-cultural activities are likely to have evolved.     

Disturbance and the structure of coral reef fish communities on the reef slope

Three levels of physical disturbance were applied to corals in permanent 10x10 m quadrats along a section of fringing reef at Lizard Island on the Great Barrier Reef to investigate the response of fish assemblages. Tabular and corymbose corals were overturned and left in situ, reducing total hard coral cover from ˜55% to ˜47%, ˜43%, and ˜34%. Despite pre-existing associations with benthic cover, all fish groups examined (pomacentrids, labrids, chaetodontids, and acanthurids) were resistent to benthic disturbances at the level and scale at which they were applied. Partial Mantel's tests, in combination with partial Canonical Correspondence Analysis enabled spatial and temporal variation to be factored out from experimental effects. Most of the variation in the fish community could be assigned to spatio-temporal variables, indicating that spatial structure over the reef landscape may moderate localised disturbance effects. This study indicates that coral reef fish assemblages may be more resistant to disturbance than many correlative studies would suggest, and highlights a need for further information on levels and scales of natural habitat disturbance in order to apply a structured approach to the experimental investigation of the importance of habitat in structuring coral-reef fish assemblages.     

Stability of a coral reef fish community following a catastrophic storm

In January 1980, a severe 3 day storm struck the normally protected leeward coral reefs of Kona, Hawaii. Waves generated by the storm, which was estimated to be a once in 20–40 year occurrence, were in excess of 6 m and caused extensive reef destruction and shoreline alteration. Shallow nearshore areas were denuded of most bottom cover and marine life. Damage to corals was extensive with broken colonies of the coral Porites compressa occurring down to depths of 27 m. Pronounced algal blooms occurred in a clearly defined sequence subsequent to the storm. The patterns of both coral destruction and algal succession were similar to those described on other storm damaged reefs. Fish mortality directly attributable to the storm was slight and the few dead fishes noted were either surge zone or tide pool species. After the storm the shallow reef flat was devoid of resident fishes while deeper areas contained many fishes which had moved from shallower water. This habitat shift substantially reduced the immediate impact of the storm on the fish community. Despite considerable habitat destruction that resulted from the storm, there were no decreases in species or population abundances on five 25 m² quadrats monitored for 23 months before and 16 months after the storm. Similarly, the numbers of triggerfish sheltering within an area increased even though the storm reduced the number of available shelter holes. The shelter and space-related carrying capacity of these areas prior to the storm may therefore not have been reached. Recolonization of evacuated areas by fishes began shortly after the storm and within 16 months many areas had regained their prestorm appearances. Large numbers of individuals remained however, in their shifted locations.     

Diurnal variability in a fish assemblage of a Bahamian coral reef

Synopsis Variation in the diurnal composition of a fish assemblage of a Bahamian coral reef was investigated by comparing visual counts of fishes taken along a 100 × 4 m wide fixed transect at four times: 0900, 1200, 1500 and 1800 hours during the summer of 1979. One sample per day was obtained at each time period over a span of 20 consecutive days. Forty-two species were recorded in these samples, with 25 occurring frequently enough to permit statistical analysis of diurnal variations in abundance. Of the 25 species compared, nearly one fourth (6 species) displayed significant variation in abundance patterns among the four time periods tested. It is suggested, because of the strong possibility of bias which might otherwise be introduced because of these variations, that repetitive quantitative visual censusing of coral reef fishes be undertaken at about the same time each day.