Damselfish

Abstract: The Great Barrier Reef (GBR) is a continental archipelagic system of 5000 reefs and shoals stretching >2000 km along the east Australia coast. The interconnectivity of these reefs should determine the choice of biological management units, which for most biota will reflect the dispersal of their eggs and/or larvae. A comparative approach using population genetics was used to ask whether the along-shore dispersal of coral reef fishes is influenced by the duration of this mobile phase. Seven species of coral reef fish, selected from three families to provide a range of taxonomic diversity and pelagic larval durations, were tested for genetic homogeneity between two regions of the GBR separated by 1000 km. A spectrum of potential dispersal capabilities was analyzed from that of Acanthochromis polyacanthus, a damselfish with brood care that uniquely lacks pelagic larvae, to that of Ctenochaetus striatus, a surgeonfish with large, specialized larvae that spend several months in the plankton. A total of 19 enzyme systems and general proteins were examined from multiple populations in each region to provide a base of 32 loci for these comparisons. With one exception, species sampled from different coral reefs within regions showed statistically significant heterogeneities across multiple loci, indicative of chaotic genetic patchiness among the samples. The exception was an anemonefish, Amphiprion melanopus, that had to be collected from large areas on each reef because of its low densities. The homogeneity of allele frequencies at local scales for this species suggests that the genetic patchiness observed in others may be a within-reef phenomenon that was manifested at the reef scale by our pseudoreplicated sampling strategy. After pooling local variability, all but two species showed significant regional differ- ences. The exceptions were the pair (Ctenochaetus striatus, Pterocaesio chrysozona) with the longest larval durations. Acanthochromis polyacanthus showed increased variation at this larger scale, consistent with a major stock division between the two regions. The logarithm of genetic variation between northern and southern populations (measured by Weir and Cockerham's Fst)was correlated with mean larval duration by an inverse linear relationship that explained 85% of the variance in the global data set. Comparison with an outgroup (Amphiprion melanopus from the Chesterfield Reefs, 1000 km east in the Coral Sea) confirmed the genetic cohesion of mainland populations for the species with shortest larval duration and shows that our empirical relationship applies only within the context of the highly connected GBR. On this basis, calculations of gene flow (Nem, the number of effective migrants per generation) between geographic regions predict panmixis for species with larval durations exceeding 1 mo. Many common species have shorter dispersal times, from which classical "isolation-by-distance" models predict differentiation between northern and southern pop- ulations at genetic equilibrium. Given that modern populations on the GBR are <10000 yr old, however, there has not been sufficient time for such differences to evolve in situ and we consider alternative scenarios for the observed heterogeneities. Comparisons with invertebrate taxa sampled over the same spatial scales imply lower gene flows in fish despite longer pelagic durations. This suggests that fish larvae may use their greater mobility to retard, rather than enhance, dispersal due to hydrodynamic advection.

Abstract: A long-term, large-scale empirical test of the recruitment limitation hypothesis was done by sampling fish populations from the southern Great Barrier Reef after having monitored their recruitment histories for 9 years. After adjustment for demographic differences, recruitment patterns explained over 90 percent of the spatial variation in abundance of a common damselfish among seven coral reefs. The age structures from individual reefs also preserved major temporal variations in the recruitment signal over at least 10 years. Abundance and demography of this small fish at these spatial and temporal scales can be explained almost entirely as variable recruitment interacting with density-independent mortality.

Abstract: The long-standing interest in density dependence in demographic rates of organisms stems from its influence in bounding population fluctuations and in shaping spatial patterns of abundance. Despite growing evidence that early mortality of marine reef fishes can be density dependent and can involve predation, the underlying biological mech- anisms have not as yet been fully explored in any system. Here we examine the causes of density-dependent juvenile mortality for two tropical damselfishes, Dascyllus flavicaudus and D. trimaculatus. These species shelter in branching corals or anemones, and they feed on plankton above their microhabitats during the day. Field experiments confirmed that density-dependent juvenile mortality of both Dascyllus species arose from predation and that most of the density-dependent loss could be attributed to small-bodied, resident piscivores (e.g., sandperch, squirrelfish) rather than larger, tran- sient species (e.g., jacks). Over the diel cycle, mortality was strongly density dependent during the dark when damselfish were sheltering but not during daylight when fish were actively foraging. Infrared video recordings revealed the species of predators responsible for most losses and indicated that most predatory events occurred from late twilight to early night, when damselfishes were in shelters and not feeding. Individuals were most at risk when located near or just outside the perimeter of a shelter. The proportion of a cohort in the riskiest areas of a microhabitat increased with density. The cause of the increased fraction of individuals at risk with increasing density was intraspecific interactions among sheltering fish jostling for space in the safest regions; this resulted in the displacement of less aggressive individuals to riskier locations. Thus, density-dependent mortality in both damselfishes arose from interference competition for refuge space from crepuscular and nocturnal predators.

Abstract: The relative roles of competition and predation in demographic density dependence are poorly known. A tractable experimental design to determine such effects and their interactions for demersal (seafloor oriented) fishes and similar sedentary species is cross-factoring multiple densities of new recruits with the presence and absence of predators. This design allows one to distinguish between density-dependent mortality due to competition alone, predation alone, or an interaction between the two, especially when supplemental field observations are available. To date, 14 species of marine fish have been examined with some variant of this design, and for 12 species predation was demonstrated to be the sole or major cause of density dependence. However, as competition may be slow acting relative to predation, the importance of competition can be underestimated in short-term experiments. On the Great Barrier Reef, we conducted a long-term field experiment in which multiple densities of new recruits of a planktivorous damselfish were cross-factored with the presence or absence of resident piscivorous fish on patch reefs. During the first 10 months, no density-dependent mortality was detected, regardless of whether resident predators were present or absent. By the end of the experiment at 17 months, per capita mortality was strongly density dependent and highly compensatory in both predator treatments; all reefs ultimately supported nearly the same adult density regardless of experimental treatment. Examination of treatment effect sizes suggested that competition was the main source of density-dependent mortality, with predation being merely a proximate agent of death. We hypothesize that predators were ineffective in this system compared with similar studies elsewhere because prey density was low relative to ample prey refuges provided by highly complex corals. Combined with previous studies, these findings indicate that density-dependent mortality in demersal marine fishes is often caused by interplay of predation and competition, whose roles may be altered by variation in habitat complexity and larval supply. These conclusions are relevant to marine fisheries models, which typically assume that density dependence is due solely to intraspecific competition.