Experimental assessment of sensory modalities of coral-reef fish larvae in the recognition of their settlement habitat

One of the great mysteries of coral-reef fish ecology is how larvae locate the relatively rare patches of coral-reef habitat on which they settle. The present study aimed to estimate, by experiments in aquaria, the sensory modalities of coral-reef fish larvae for senses used in searching for their species settlement habitat. Larval recognition of settlement habitat can be based on the detection of conspecifics and/or of characteristics of coral habitat using visual, chemical and mechanical cues. For this study, larvae were captured with crest nets and were then introduced into experimental tanks that allowed testing of each type of cue separately (visual, chemical or mechanical cues). Among the 18 species studied, 13 chose their settlement habitat due to the presence of conspecifics and not based on the characteristics of coral habitat, and 5 species did not move toward their settlement habitat (e.g. Scorpaenodes parvipinnis, Apogon novemfasciatus). Among the different sensory cues tested, two species used the three types of cues (Parupeneus barberinus and Ctenochaetus striatus: visual, chemical and mechanical cues), six used two types (e.g. Myripristis pralinia: visual and chemical cues; Naso unicornis: visual and mechanical cues), and five used one type (e.g. Chrysiptera leucopoma: visual cues; Pomacentrus pavo: chemical cues). These results demonstrate that many coral-reef fish larvae could in practice use sensory cues for effective habitat selection at settlement, and have the ability to discriminate species-specific sensory cues.Communicated by J. Krause     

Local variation in mortality rates and life-expectancy estimates of the coral-reef fish Centropyge bicolor (Pisces: Pomacanthidae)

Mortality rates of the coral reef fish Centropyge bicolor were based on the disappearance of known individual fish. Data were collected over three years from 1978 at four sites around Lizard Island, Queensland, Australia. To measure disappearance, membership of 36 harems (comprising about 150 fish at any one time) was determined at bimonthly intervals. Mortality rates varied significantly among the four study areas. A tenfold difference in mean mortality occurred between two of the areas. The ranking of areas according to mortality rate was similar for 1978–1979 and 1979–1980. The ranking of areas was again similar for 1980–1984 when mortality rates were determined from an additional census in June 1984. Average further life-expectancy of mature fish based on mortality rates varied from 1 yr in one study area to 5 to 13 yr in a different area.     

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.     

Effects of nano carbon black and single-layer graphene oxide on settlement,survival and swimming behaviour of Amphibalanus amphitrite larvae

The effects of two carbon-based nanomaterials, nano-sized carbon black (nCB), and single-layer graphene oxide (GO) on settlement of Amphibalanus amphitrite (Cirripedia, Crustacea) cypris larvae (cyprids) were assessed after 24, 48, and 72 h of exposure. Additionally, the effects of these nanomaterials on the mortality and swimming behaviour of the nauplius larvae (nauplii) of the same organism were determined after 24 and 48 h. The data indicate that nCB is more effective as a potential antisettlement agent than single-layer GO; moreover, nCB did not show any adverse effects on the larvae. The swimming behaviour of II stage nauplii of A. amphitrite exposed to a suspension of nCB was inhibited only at very high nCB concentrations (≥0.5 mg/mL). Single-layer GO, on the contrary, showed lower antisettlement effects and was more active in altering the survival and inhibiting the swimming behaviour of the nauplii. An indication of the toxic or non-toxic mechanisms of the antisettlement properties of both of these nanomaterials is provided by the reversibility of the antisettlement activity. In conclusion, we propose nCB as an innovative antifouling nanomaterial that shows low toxicity towards the model organism (crustaceans) used in this study.     

Auditory and olfactory abilities of pre-settlement larvae and post-settlement juveniles of a coral reef damselfish (Pisces: Pomacentridae)

The propagules of most species of reef fish are advected from the reef, necessitating a return to reef habitats at the end of the pelagic stage. There is increasing evidence of active attraction to the reef but the sensory abilities of reef fish larvae have not been characterized well enough to fully identify cues. The electrophysiological methods of auditory brainstem response (ABR) and electroolfactogram (EOG) were used to investigate auditory and olfactory abilities of pre- and post-settlement stages of a damselfish, Pomacentrus nagasakiensis (Pisces, Pomacentridae). Audiograms of the two ontogenetic stages were similar. Pre-settlement larvae heard as well as their post-settlement counterparts at all but two of the tested frequencies between 100 Hz and 2,000 Hz. At 100 and 600 Hz, pre-settlement larvae had ABR thresholds 8 dB higher than those of post-settlement juveniles. Both stages were able to detect locally recorded reef sounds. Similarly, no difference in olfactory ability was found between the two ontogenetic stages. Both stages showed olfactory responses to conspecifics as well as L-alanine. Therefore, the auditory and olfactory senses have similar capabilities in both ontogenetic stages. Settlement stage larvae of P. nagasakiensis can hear and smell reef cues but it is unclear as to what extent larvae use these sounds or smells, or both, as cues for locating settlement sites. An erratum to this article can be found at     

Eggs and early larvae of Thrissocles species (Engraulidae: Pisces)

Eggs of Thrissocles species are found in surface plankton in the Ernakulam Channel (Cochin Harbour)during February to May 1967. The eggs hatch within 24 h. Empty egg shells have characteristic apertures, through which the embryos have emerged; yolk is resorbed 36 h after hatching.Larvae (36 to 72 h groups) assemble at the lighted region of the aquaria during day-time and scatter to different levels at night. Larvae older than 72 h show no inclination to assemble as before. All larvae died between 96 and 110 h after hatching. Many batches of eggs were reared in the same medium, and all of them behaved as described. The results indicate that the right type of food was available in the aquaria for larvae up to a period of 72 h. The volume of water also appears to have a bearing upon the survival rate and longevity of the larvae since, in small aquaria, more larvae died at an early stage.     

In situ ontogeny of behaviour in pelagic larvae of three temperate, marine, demersal fishes

The ontogeny of behaviour relevant to dispersal was studied in situ with reared pelagic larvae of three warm temperate, marine, demersal fishes: Argyrosomus japonicus (Sciaenidae), Acanthopagrus australis and Pagrus auratus (both Sparidae). Larvae of 5–14 mm SL were released in the sea, and their swimming speed, depth and direction were observed by divers. Behaviour differed among species, and to some extent, among locations. Swimming speed increased linearly at 0.4–2.0 cm s⁻¹ per mm size, depending on species. The sciaenid was slower than the sparids by 2–6 cm s⁻¹ at any size, but uniquely, it swam faster in a sheltered bay than in the ocean. Mean speeds were 4–10 body lengths s⁻¹. At settlement size, mean speed was 5–10 cm s⁻¹, and the best performing individuals swam up to twice the mean speed. In situ swimming speed was linearly correlated (R ²=0.72) with a laboratory measure of swimming speed (critical speed): the slope of the relationship was 0.32, but due to a non-zero intercept, overall, in situ speed was 25% of critical speed. Ontogenetic vertical migrations of several metres were found in all three species: the sciaenid and one sparid descended, whereas the other sparid ascended to the surface. Overall, 74–84% of individual larvae swam in a non-random way, and the frequency of directional individuals did not change ontogenetically. Indications of ontogenetic change in orientated swimming (i.e. the direction of non-random swimming) were found in all three species, with orientated swimming having developed in the sparids by about 8 mm. One sparid swam W (towards shore) when <10 mm, and changed direction towards NE (parallel to shore) when >10 mm. These results are consistent with limited in situ observations of settlement-stage wild larvae of the two sparids. In situ, larvae of these three species have swimming, depth determination and orientation behaviour sufficiently well developed to substantially influence dispersal trajectories for most of their pelagic period.     

Food selection by laboratory-reared larvae of the scaled sardine Harengula pensacolae (Pisces,Clupeidae) and the bay anchovy Anchoa mitchilli (Pisces,Engraulidae)

Food selection by laboratory-reared larvae of scaled sardines Harengula pensacolae, and bay anchovies Anchoa mitchilli, was compared. Natural plankton was fed to the larvae during the 22 days following hatching. Food levels in the rearing tanks were maintained at an average of 1,600 to 1,800 potential food organisms per liter. Larvae of both species selected as food copepod nauplii, copepodites, and copepods; initial feeding was on organisms of 50 to 75 body width. Larvae of H. pensacolae averaged 4.2 mm in total length at hatching and those of A. mitchilli about 2.0 mm. H. pensacolae larvae grew about 1.0 mm per day and A. mitchilli 0.70 mm per day. The mean number of food organisms in each digestive tract was greater in H. pensacolae than in A. mitchilli, and the difference in number increased as the larvae grew. Average size of food organisms eaten increased for both species with growth, because of selection by the larvae; the average size of copepodites and copepods in digestive tracts increased at a faste rate in H. pensacolae than A. mitchilli. A. mitchilli longer than 8 mm did not eat copepod nauplii.Contribution No. 170, Bureau of Commercial Fisheries Tropical Atlantic Biological laboratory, Miami, Florida 33149, USA.     

Partitioning behavior and intra- and interspecific interactions: a comparison between male and female groupers,Cephalopholis spiloparaea (Pisces: Serranidae: Epinephelinae)

Behavior of male and female groupers, Cephalopholis spiloparaea, observed, between 1990 and 1991 in Sasanhaya Bay, Rota, Northern Mariana Islands, was compared to detect differences in proportional use between sexes, and between daylight and pre-courtship time periods. This species has male-dominated haremic groups. Time partitioning could not be measured exactly. Instead, the numbers of intra-and interspecific interactions, foraging, resting, swimming, and maintenance behaviors of each sex were measured during the observation periods. The proportion of acts in each category out of the total number of acts, termed effort, was used in comparisons. Males were predicted to devote more effort toward intra-and interspecific interactions compared to females and to maximize reproductive success. Females were predicted to devote more effort towards foraging, compared to males. This behavior maximizes reproductive effort. Both were predicted to engage in greater intra-and interspecific interactions during the pre-courtship period, corresponding both to the approaching time of courtship and to increases in the numbers of potential spatial and trophic competitors. Intra-and interspecific interactions of both males and females were greater during the pre-courtship period. Differences in all classes of behavior between sexes were only weakly significant for two, however. Males had greater effort in intra-and interspecific interactions. Females had greater effort only in maintenance behavior. Both differences were evident only during the pre-courtship period. Foraging behavior by both sexes was virtually absent during daylight and pre-courtship periods. Fish sought shelter and were not incidentally observed foraging after dusk, suggesting that this species actively forages later at night or during early morning hours, just prior to and during sunrise.     

Right tools for the task: intraspecific modality in the swimming behaviour of coral reef fishes

Understanding the movement patterns of fishes in the wild requires an awareness of the intrinsic and extrinsic factors underlying their behaviour. Using field observations, we explored whether eight species of coral reef fish display modality in their swimming behaviour, which is a convergence in fin use patterns among and within species when they conduct daily activities (e.g. feeding, travelling, chasing). While a range of different fins were used during steady swimming by each species, most fishes converged towards median-paired fin use (e.g. pelvics, pectorals) when undertaking searching and feeding. Conversely, all species adopted caudal fin use during high-speed activities such as chasing. Such modality in fin use was relatively conserved across juvenile and adult conspecifics and transcended differences in phylogenetic history and trophic preferences, suggesting that overarching constraints underpin patterns of fish swimming behaviour in complex habitats.     

Environmental influences on larval duration,growth and magnitude of settlement of a coral reef fish

The influence of environmental variables on the planktonic growth, pelagic larval duration and settlement magnitude was examined for the coral reef surgeonfish Acanthurus chirurgus. Newly settled fish were collected daily from patch reefs in the San Blas Archipelago, Caribbean Panama for 3.5 years. Environmental influences on growth were examined at three different life history stages: from 0 to 6 days, 7 to 25 days and from 26 to 50 days after hatching. Larval growth was correlated, using multiple regression techniques, with a combination of factors including solar radiation, rainfall, and along-shore winds. Depending on the life history stage, these accounted for 13–38% of the variation in growth rates when all the months were included in the analyses. Correlations between environmental variables and growth also varied among seasons and were stronger in the dry than in the wet season. During the dry season solar radiation, rainfall and along-shore winds described 57%, 86% and 74% of the variability in growth between 0 and 6 days, 7 and 25 days and 26 and 50 days, respectively. During the wet season rainfall, along-shore winds and temperature only described 38% of the variability in early growth and 27% of growth just before settlement. No significant model was found to describe growth 7–25 days after hatching during the wet season. Rainfall, solar radiation and along-shore winds were negatively correlated with growth up to 25 days after hatching but positively correlated as larvae approached settlement at a mean age of 52 days. Over 65% of the variability in pelagic larval duration was accounted for by a regression model that included solar radiation and along-shore winds. When data sets from wet and dry seasons were analysed separately, along-shore winds accounted for 67% of the change in larval duration in the dry season, and solar radiation accounted for 23% of the variation in larval duration in the wet season. Only 22% of the variability in settlement intensity could be described by solar radiation and temperature, when all months of the year were included in the analysis. Solar radiation and rainfall were included in a regression model that accounted for 40% of the variation in numbers of fish settling during the dry season. This study suggests that the levels of solar radiation, along-shore winds and rainfall during the early larval life can have important effects on the growth, larval duration and consequently, the settlement magnitude of marine fishes. Results also highlight the need to account for seasonality and ontogeny in studies of environmental influences.Communicated by G.F. Humphrey, Sydney     

Infestation dynamics of gnathiid isopod juveniles parasitic on the coral-reef fish Hemigymnus melapterus (Labridae)

The dynamics of infestation by parasitic juveniles of gnathiid isopods were investigated at Lizard Island in the summer of 1997/1998 to determine when, and at what rate, they infest fishes. Variation in gnathiid abundance on wild-caught fish (Hemigymnus melapterus) between dawn and sunset was examined, and unparasitized H. melapterus in cages were exposed to gnathiids in the field for 4 h (8 h for fish sampled at 06:00 h) at five different times of the day and night (10:00, 14:00, 18:00, 22:00, 06:00 hrs) on three reefs. To control for any potential effect of the cleaner-fish Labroides dimidiatus, which consumes large numbers of gnathiids each day, sampling was carried out on reefs from which all cleaner-fish had been removed. Gnathiid abundance decreased during the day. Standardized abundance per wild-caught fish was 1.9 times higher at dawn than at sunset. Gnathiids successfully infested fish in cages. Sixty-one percent of the fish in cages were infested with ≥1 gnathiids, with 51% of the fish having between 1 and 4 gnathiids after 4 h exposure. Gnathiids infested fish both day and night. The log₁₀ (abundance +1) of gnathiids per caged fish varied significantly between time periods, with higher gnathiid numbers on fish sampled at 18:00, 22:00, and 06:00 hrs than on fish sampled at 10:00 and 14:00 hrs. In contrast to gnathiid abundance on wild-caught fish, the number of gnathiids on caged fish sampled at sunset was as high as that at dawn. The estimated mean (SE) cumulative number of gnathiids per caged fish [mean size = 13.7 cm (±0.25)] per day was 7.8 (1.1); this is similar to the estimated mean number of 7.3 gnathiids on similar-sized wild-caught fish at 6:00 hrs. The high infestation rate of gnathiids on caged fish in the late afternoon contrasted with the low numbers on wild-caught fish at this time, suggesting that factors other than infestation behaviour may be responsible for the low numbers of gnathiids on wild-caught fish in the afternoon. Received: 19 August 1998 / Accepted: 1 June 1999     

Experimental analysis of the contribution of swimming and drifting to the displacement of reef fish larvae

The extent to which behaviour affects the dispersal of pelagic larvae in reef fishes has been a topic of major discussion among marine ecologists. Here, we experimentally quantified the extent to which the displacement of late-stage larvae of Abudefduf saxatilis is due to active movement (i.e. swimming) and drifting. We consider drifting as the component of larval displacement accounted for by the current. Drifting was quantified by comparing larval displacement to the displacement of passive particles in an extended flow chamber that gave larvae the free choice of swimming (i.e. swim with or against the current or not swim at all). We also determine whether drifting results from currents exceeding larval swimming capabilities or from the behavioural choice of larvae of not to swim against adverse currents. To do this, we compare the speeds of larval swimming in the extended flow chamber to those obtained in a smaller chamber in which larvae are behaviourally forced to swim due to space constraints and a retaining fence (most available data on larval swimming is based on this sort of chamber). Within the extended chamber, larvae tended to face the current and swim slower than it. This resulted in a net displacement increasingly determined by drifting. We also found that in the extended chamber, larvae swam at speeds between one and six times slower than the speeds they achieved in the “behaviourally modifying” smaller chamber. This suggests that the net displacement in the extended chamber was in part due to the behavioural choice of the larvae of not to swim. The importance of this “behavioural drifting” is discussed in terms of energy savings required for successful completion of the larval period and post-settlement survival. The idea that larvae may modulate their swimming behaviour raises caution for the use of published data regarding swimming capabilities of reef fish larvae when assessing the extent to which these fish actively affect their dispersal.     

On the evolution of nest raiding and male defensive behaviour in sticklebacks (Pisces: Gasterosteidae)

Summary We develop a model to explain the evolution of nest raiding by female sticklebacks based on enhanced female reproductive success derived by creating mating opportunities with the male whose nest has been destroyed. Our model depends critically on the shape of the function relating percentage hatch success to the number of eggs in a male's nest, and the probability that the female will later mate with the male whose nest was raided. Raiding by females will be favoured when the number of eggs in the nest is high providing high egg numbers reduces the percentage hatch rate. High probabilities of mating with the male in question also favour raiding. A second model suggests that males cannot reduce the probability of being raided by limiting the number of eggs in the nest. Instead selection should favour their increasing the number of eggs in the nest. Tests of these models are suggested and their relation to other factors which may influence the evolution of cannibalism are discussed.     

Vertical and horizontal distribution of fish larvae near coral reefs at Lizard Island,Great Barrier Reef

Consistent patterns of horizontal distribution of fish larvae from plankton tows were found in shallow waters around Lizard Island, Great Barrier Reef during 1979 and 1980. Few types of larvae were most abundant in Lizard Lagoon, and none of these were old larvae. Forty percent of the 57 types of larvae studied differed in abundance between windward and downwind sides of the island. More types of old larvae were found in greatest abundance off the windward side of the island than the downwind side. Most types of larvae preferred deeper water (>3 m) during the day and moved upward at night, although a few types preferred upper (<3 m) or middle portions of the water column. These latter were more likely to descend at night or to maintain their day-time distribution than to move upward. Windward larvae [those more abundant off the windward (SE) side of the island] were more shallow-living than were downwind larvae, and were more likely to maintain their day-time distribution at night. The current patterns around Lizard Island were favourable for retention of larvae in both Lizard Lagoon and off the windward side of the island, if combined with certain vertical distributions of the larvae. However, while there was evidence for retention on the windward side of the island, there was no evidence for retention in Lizard Lagoon. Currents on the downwind side of the island were not favourable for retention of larvae and there was little evidence that larvae were retained there. Retention may be an accidental result of interaction between currents and larval behaviour, or the result of a strategy of retention by the larvae. These could not be distinguished in the present study.     

The ecology of Indo-Pacific grouper (Serranidae) species and the effects of a small scale no take area on grouper assemblage,abundance and size frequency distribution

This paper used the case study of the Wakatobi Marine National Park, Indonesia to examine changes in the diversity, density and maturity of grouper species over a 5-year period following the establishment of a small-scale no-take area (NTA). This work was carried out to investigate whether “small” NTAs could be effective management strategies over a time scale that is relevant to local fishery communities and their perception of management success. Our research also documents the ecology of these species, information essential if we are to understand how management practises are to affect coral reef fish species. Designation of this “small NTA” increased the density of groupers by 30% over a 5-year period of protected status. After 5 years of protection, grouper populations within this NTA were more mature and double the density of those within the adjacent lightly fished sites and nearly five times those of a heavily fished site. During this time all other nearby fished sites underwent large declines in grouper density. The nearby lightly fished Kaledupa site decreased by up to 50% year⁻¹. Such drastic declines are considered the impact of the exponential development of ever efficient and unsustainable methods of fishing within the study region. This NTA was not of benefit to all grouper species; the reasons for which are not clear. Such questions require further detailed research about the life history, population and behavioural ecology of Indo-Pacific grouper species. Such information is critical for urgently needed fisheries management. The present study found that a small scale NTA of 500 m length was large enough to increase the population of top predatory fish. In conjunction with other socially acceptable small scale NTAs it could help maintain and increase important fish stocks over a larger area. The use of “small” NTAs within networks of reserves should become a useful tool in the management of the locally exploited coral reefs.     

Effects of temperature on oxygen consumption,growth, and development of embryos and yolk-sac larvae of Siganus randalli (Pisces: Siganidae)

Eggs from laboratory spawnings of the coralreef fish Siganus randalli Woodland were incubated at two temperatures (27 and 30 °C). Eggs and larvae were sampled until larval starvation, while changes in oxygen consumption, growth, yolk utilization, and development were monitored. Oxygen consumption, which peaked at hatching, was higher for embryos incubated at 30 °C than at 27 °C. Rates of oxygen consumption (nl h^-1 individual^-1) at hatching were similar to those for other temperate and tropical species. Rates of oxygen consumption by yolk-sac larvae were highly variable, and these data suggest that larval oxygen consumption prior to yolk-sac absorption may not be significantly influenced by temperature. Rates of yolk depletion were higher for larvae at the higher temperature. After an initial rapid increase in length, length of larvae at 30 °C decreased with age. Egg size, egg weight, and maximum notochord length of larvae differed significantly between spawns. Age-specific oxygen consumption rates by the embryos varied between spawns, but regressions describing oxygen consumption as a function of age did not differ significantly. The initiation and completion of eye pigmentation were used as developmental markers to calculate the amount of yolk remaining for larvae at the different temperatures. Larvae maintained at 30 °C completed eye pigmentation approximately 3 h sooner than those maintained at 27 °C, but had less endogenous reserves. This finding indicates a trade-off between rapid development and efficient utilization of the endogenous reserves. The completion of eye pigmentation in larvae incubated at the higher temperature occurred at midnight and, depending on the amount of time that the larvae have to initiate feeding prior to the point-of-no-return, the timing of completion of eye pigmentation could influence larval survival.