Resource limitation and recruitment patterns in a coral reef fish assemblage

The potential effects of food and shelter availability on the recruitment and early survivorship of coral reef fishes were studied on St. Croix, U.S. Virgin Islands. The faunal assemblage studied included diurnally active fishes found in the “rubble/sand” habitat. The most abundant members were: beaugregory, Stegastes leucostictus (Muller & Troschel), goldspotted goby, Gnatholepis thompsoni Jordan, bridled goby, Coryphopterus glaucofraenum Gill, surgeonfishes, Acanthurus bahianus Castelnau and A. chirurgus (Bloch), and French grunt, Haemulon flavolineatum (Desmarest). Comparisons of recruitment to reefs constructed from substrata that varied in morphological characteristics showed that there were differences in the relative abundances of recruits attracted to and/or surviving on the different reef types. Juveniles of most species appeared to prefer the branching coral Porites porites (Pallas), which provided a large number of small crevices between the branches.Manipulations of the availability of shelter sites for fishes demonstrated that recruitment and/or early survivorship were strongly limited by the number of refuges. This result was found in six separate carried out during different years and in different seasons. Shelter site availability presumably limits fish populations through its effects on prédation rates.Experimental manipulations of food availability indicated that food does not directly influence settlement or early survivorship of coral reef fishes. However, it is probable that correlations between habitat characteristics and food availability have influenced the evolution of settling preferences.     

Communication in coral reef fish: the role of ultraviolet colour patterns in damselfish territorial behaviour

Many coral reef fish possess ultraviolet (UV) colour patterns. The behavioural significance of these patterns is poorly understood and experiments on this issue have not been reported for free-living reef fish in their natural environment. The damselfish Pomacentrus amboinensis has UV facial patterns, and spectroradiometric ocular media measurements show that it has the potential for UV vision. To test the potential behavioural significance of the UV patterns, I studied the response of males, in natural territories on the reef and in aquaria, to two conspecific intruders, one presented in a UV-transmitting (UV+) container and the other in a UV-absorbing (UV−) one. Territory owners attacked intruders viewed through UV+ filters significantly more often and for longer than intruders viewed through the UV− filter. In general, the results of the field experiment confirmed those of the laboratory experiment. The results support the hypothesis that P. amboinensis males are sensitive to UV light and that reflectance patterns, which appear in high contrast only in UV, modulate the level of aggressive behaviour. A recent survey showed that many predatory fish may not have UV vision and the use of UV colours in select species of reef fish may therefore serve as a ‘private communication channel’.     

Density-associated recruitment mediates coral population dynamics on a coral reef

Theory suggests that density-associated processes can modulate community resilience following declines in population size. Here, we demonstrate density-associated processes in two scleractinian populations on the outer reef of Moorea, French Polynesia, that are rapidly increasing in size following the effects of two catastrophic disturbances. Between 2006 and 2010, predation by the corallivorous crown-of-thorns sea star reduced coral cover by 93 %; in 2010, the dead coral skeletons were removed by a cyclone, and in 2011 and 2012, high coral recruitment initiated population recovery. Coral recruitment was associated with coral cover, but the relationship differed between two coral genera that are almost exclusively broadcast spawners in Moorea. Acroporids recruited at low densities, and the density of recruits was positively associated with cover of Acropora, whereas pocilloporids recruited at high densities, and densities of their recruits were negatively associated with cover of Pocillopora. Together, our results suggest that associations between adult cover and density of both juveniles and recruits can mediate rapid coral community recovery after large disturbances. The difference between taxa in sign of the relationships between recruit density and coral cover indicate that they reflect contrasting mechanisms with the potential to mediate temporal shifts in taxonomic composition of coral communities.     

Evidence of density-independent mortality in a settling coral reef damselfish, Chromis viridis

To know if the variation in the number of settling fish larvae can be dampened by density-dependent postsettlement mortality, we investigated the relationship between settler density and predator-induced mortality of a coral reef damselfish, Chromis viridis. Totals of 2, 3, 5, 8, 10, 12, 14, 16, 18, and 20 fish of 10 or 20 mm total length were released in experimental cages enclosing a coral head of Porites rus (to provide settlement habitat) and five predators. The results showed that the mortality rate of both 10- and 20-mm fish was density independent.     

Effects of territorial damselfish on an algal-dominated coastal coral reef

Territorial damselfish are important herbivores on coral reefs because they can occupy a large proportion of the substratum and modify the benthic community to promote the cover of food algae. However, on coastal coral reefs damselfish occupy habitats that are often dominated by unpalatable macroalgae. The aim of this study was to examine whether damselfish can maintain distinctive algal assemblages on a coastal reef that is seasonally dominated by Sargassum (Magnetic Island, Great Barrier Reef). Here, three abundant species (Pomacentrus tripunctatus, P. wardi and Stegastes apicalis) occupied up to 60% of the reef substrata. All three species promoted the abundance of food algae in their territories. The magnitudes of the effects varied among reef zones, but patterns were relatively stable over time. Damselfish appear to readily co-exist with large unpalatable macroalgae as they can use it as a substratum for promoting the growth of palatable epiphytes. Damselfish territories represent patches of increased epiphyte load on macroalgae, decreased sediment cover, and enhanced cover of palatable algal turf.     

El Niño mass coral mortality: a test of resource limitation in a coral reef damselfish population

Summary The mass mortality of reef corals in the eastern Pacific as a result of the 1982–1983 El Niño oceanographic anomaly permitted the first large scale test of resource limitation for a coral reef fish. Population densities of territorial herbivorous damselfish did not respond to the massive regional increase in space available for the cultivation of algal food following the El Niño event. The proportion of juveniles in the population was low and new recruits were uncommon, indicating that recruitment rates rather than resource supply probably control the abundance of this coral reef damselfish.     

Nonindependent mating in a coral reef damselfish: evidence of mate choice copying in the wild

Theoretical and experimental studies have shown that mate choicecopying is a viable mating strategy under certain conditions.Copying experiments in fish have been conducted primarily inthe laboratory, except for one study conducted in the fieldunder artificial conditions. We investigated whether in a wildpopulation of the coral reef whitebelly damselfish (Amblyglyphidodonleucogaster) females copy the choice of other females. Femalespreferentially spawn with males that have recently mated. Todetermine if the presence of new eggs in the nest was the reasonfemales chose mates or whether females were mate choice copying,we conducted egg-switching experiments. Eggs from males thatrecently mated were donated to males that had no eggs. If femalesare mate choice copying, then donor males with no eggs in thenest should continue to receive additional eggs. If femalesare using the presence of new eggs as the criterion for matechoice, then foster males with new eggs should receive additionaleggs. We found that donor males received new eggs significantlymore often than expected. More females mated with donor malesthan foster males. Furthermore, females preferentially choseto mate with males whom they had seen mating with another female.Females appear to remember the mate choice of other femalesand choose to mate with those same males even after 1 day. Theseresults suggest that females may be copying the mating decisionof other females rather than choosing males based on the presenceof new eggs in the nest.     

Patterns of recruitment of coral reef fishes in a monsoonal environment

The abundance of newly settled recruits of coral reef fishes was monitored at a total of 11 sites at two islands and two coastal locations in the central Philippines for a 20-month period (February 2008 to September 2009) that included two monsoon cycles. Recruitment occurred throughout the year. Most of the abundant species exhibited protracted recruitment seasons. This confirms the expectation of extended breeding of reef fishes at lower latitudes. The annual pattern of recruitment of reef fishes as a group was predictable. Annual fluctuations of sea surface temperature and wind strength largely explained the pattern. Rainfall, however, did not significantly influence the pattern of recruitment. Peaks in density and species richness of recruits occurred during the southwest monsoon and the second inter-monsoonal period of the year (July to October) when temperatures were highest and when most of the sites were sheltered from winds or when winds were weak. Conversely, lowest density and species richness were observed during the northeast monsoon (November to March) when temperatures were lowest and most sites were exposed to winds. The same pattern could also be seen in the recruitment of both damselfishes (Pomacentridae) and wrasses (Labridae), notwithstanding a tenfold difference in abundance of recruits between the two families. The pattern was fairly consistent across most sites, among most of the species that were examined, and between the 2 years that were sampled. This study is one of the few to provide insights into the influence of environmental factors on the recruitment patterns of fishes on Indo-Pacific coral reefs situated at lower latitudes.     

Emergent patterns of population genetic structure for a coral reef community

What shapes variation in genetic structure within a community of codistributed species is a central but difficult question for the field of population genetics. With a focus on the isolated coral reef ecosystem of the Hawaiian Archipelago, we assessed how life history traits influence population genetic structure for 35 reef animals. Despite the archipelago's stepping stone configuration, isolation by distance was the least common type of genetic structure, detected in four species. Regional structuring (i.e. division of sites into genetically and spatially distinct regions) was most common, detected in 20 species and nearly in all endemics and habitat specialists. Seven species displayed chaotic (spatially unordered) structuring, and all were nonendemic generalist species. Chaotic structure also associated with relatively high global F_ST. Pelagic larval duration (PLD) was not a strong predictor of variation in population structure (R² = 0.22), but accounting for higher F_ST values of chaotic and invertebrate species, compared to regionally structured and fish species, doubled the power of PLD to explain variation in global F_ST (adjusted R² = 0.50). Multivariate correlation of eight species traits to six genetic traits highlighted dispersal ability, taxonomy (i.e. fish vs. invertebrate) and habitat specialization as strongest influences on genetics, but otherwise left much variation in genetic traits unexplained. Considering that the study design controlled for many sampling and geographical factors, the extreme interspecific variation in spatial genetic patterns observed for Hawaìi marine species may be generated by demographic variability due to species‐specific abundance and migration patterns and/or seascape and historical factors.     

Habitat selection and aggression as determinants of spatial segregation among damselfish on a coral reef

Adults of many closely related coral reef fish species are segregated along gradients of depth or habitat structure. Both habitat selection by new settlers and subsequent competitive interactions can potentially produce such patterns, but their relative importance is unclear. This study examines the potential roles of habitat selection and aggression in determining the spatial distribution of adults and juveniles of four highly aggressive damselfishes at Lizard Island, northern Great Barrier Reef. Dischistodus perspicillatus, D. prosopotaenia, D. melanotus, and D. pseudochrysopoecilus maintain almost non-overlapping distributions across reef zones, with adults of one species dominating each reef zone. Juveniles exhibit slightly broader distributional patterns suggesting that subsequent interactions reduce overlap among species. Although habitat choice experiments in aquaria suggest that associations between juveniles and substrata types in the field are partly due to habitat selection, large overlaps in the use of substrata by the different species were also found, suggesting that substratum selection alone is insufficient in explaining the discrete spatial distributions of adults. The strength of aggressive interactions among all four species was tested by a "bottle" experiment, in which an adult or juvenile of each species was placed in the territories of adult fish on the reef. The greatest levels of interspecific aggression were directed against adults and juveniles of neighbouring species. The highest levels of aggression were associated with species exhibiting the greatest levels of overlap in resource use. Evidently both habitat selection and interspecific aggression combine to determine the adult distributions of these species.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

Habitat choice,recruitment and the response of coral reef fishes to coral degradation

The global degradation of coral reefs is having profound effects on the structure and species richness of associated reef fish assemblages. Historically, variation in the composition of fish communities has largely been attributed to factors affecting settlement of reef fish larvae. However, the mechanisms that determine how fish settlers respond to different stages of coral stress and the extent of coral loss on fish settlement are poorly understood. Here, we examined the effects of habitat degradation on fish settlement using a two-stage experimental approach. First, we employed laboratory choice experiments to test how settlers responded to early and terminal stages of coral degradation. We then quantified the settlement response of the whole reef fish assemblage in a field perturbation experiment. The laboratory choice experiments tested how juveniles from nine common Indo-Pacific fishes chose among live colonies, partially degraded colonies, and dead colonies with recent algal growth. Many species did not distinguish between live and partially degraded colonies, suggesting settlement patterns are resilient to the early stages of declining coral health. Several species preferred live or degraded corals, and none preferred to associate with dead, algal-covered colonies. In the field experiment, fish recruitment to coral colonies was monitored before and after the introduction of a coral predator (the crown-of-thorns starfish) and compared with undisturbed control colonies. Starfish reduced live coral cover by 95–100%, causing persistent negative effects on the recruitment of coral-associated fishes. Rapid reductions in new recruit abundance, greater numbers of unoccupied colonies and a shift in the recruit community structure from one dominated by coral-associated fishes before degradation to one predominantly composed of algal-associated fish species were observed. Our results suggest that while resistant to coral stress, coral death alters the process of replenishment of coral reef fish communities.     

Spatial patterns in the distribution of damselfishes on a fringing coral reef

Damselfishes are an important element of the fauna of coral reefs. This study describes spatial patterns in the distribution of 15 species of damselfishes at Lizard Island, northern Great Barrier Reef (GBR). The aim of the work was to identify the spatial scales at which major changes in the composition and abundance of the fauna occurred. These patterns were then compared with previous studies in an attempt to determine if distributions followed general patterns at a range of localities. The assemblage found at Lizard Island was similar to that of reefs in the central GBR. The most important changes in the composition of the fauna occurred among reef zones. Shallow zones (the reef flat and crest) were dominated by herbivorous species while planktivorous and omnivorous species were most abundant in deeper zones (the reef slope). Densities of herbivorous damselfishes in shallow reef zones at Lizard Island averaged 45.5 individuals per 80 m², a value comparable to densities found in similar zones on reefs in the central and southern GBR and at one locality in the Caribbean. Comparisons of relative distributions suggested that abundant species tend to be widely distributed among zones and habitats, while rare species have restricted distributions at Lizard Island. However, computer simulation of the sampling program suggested that the ability of our study to describe the distribution patterns of rare species was limited, despite intensive sampling. Correlations between breadth of distribution and abundance may have occurred simply because rare species were less likely to be recorded within a transect. Our results suggest that it will be difficult to compare the distribution patterns of species among studies. Furthermore, the interpretation of relative patterns of distribution at a single locality in terms of ecological specialization or partitioning may first require an assessment of the ability of the sampling program to accurately record spatial patterns.     

Dispersal in the spiny damselfish,Acanthochromis polyacanthus,a coral reef fish species without a larval pelagic stage

The spiny damselfish, Acanthochromis polyacanthus, is widely distributed throughout the Indo‐Australian archipelago. However, this species lacks a larval dispersal stage and shows genetic differentiation between populations from closely spaced reefs. To investigate the dispersal strategy of this unique species, we used microsatellite markers to determine genetic relatedness at five dispersal scales: within broods of juveniles, between adults within a collection site (~30 m²), between sites on single reefs, between nearby reefs in a reef cluster, and between reef clusters. We sampled broods of juveniles and adults from seven reefs in the Capricorn‐Bunker and Swain groups of the Great Barrier Reef. We found that extra‐pair mating is rare and juveniles remain with their parents until fledged. Adults from single sites are less related than broods but more related than expected by chance. However, there is no evidence of inbreeding suggesting the existence of assortative mating and/or adult migration. Genetic differences were found between all of the reefs tested except between Heron and Sykes reefs, which are separated only by a 2‐km area of shallow water (less than 10 m). There was a strong correlation between genetic distance, geographical distance and water depth. Apparently, under present‐day conditions spiny damselfish populations are connected only between sites of shallow water, through dispersal of adults over short distances. Assuming that dispersal behaviour has not changed, the broad distribution of A. polyacanthus as a species is likely based on historical colonization patterns when reefs were connected by shallow water at times of lower sea levels.     

Food ration and condition affect early survival of the coral reef damselfish, Stegastes partitus

The supply of larvae is a major determinant of population and community structure in coral reef fishes. However, spatial and temporal variation in condition (i.e. quality) of potential recruits, as well as their density (i.e. quantity), may influence survival and growth of juveniles. We conducted an experiment to test whether recent feeding history could affect growth, condition and post-recruitment survival in a Caribbean damselfish, Stegastes partitus. Fish were collected soon after settlement, and fed either low or high rations in aquaria for 7 days. Fish fed the high ration grew faster in aquaria and were in a better condition (higher total lipids and Fulton’s condition factor) at the end of the feeding period. Subsequently, we released 50 fish in 25 pairs (one fish subjected to low rations, the other to high rations) on a Bahamian coral reef and monitored survival for 10 days. Survivorship of high-ration fish was double that of low-ration fish (80 vs 40% over 10 days). However, low-ration fish that survived 10 days were of similar condition and grew at similar rates to high-ration fish, suggesting that short-term ration differences may not persist in surviving fish. Laboratory experiments showed that low-ration fish were taken by piscivorous fishes before high-ration fish, indicating that differential predation may account for survival differences. This study highlights the potential of feeding history and condition to affect the relationship between patterns of larval arrival at reefs, and subsequent juvenile and adult population densities. Received: 1 March 1999 / Accepted: 15 July 1999     

From record performance to hypoxia tolerance: respiratory transition in damselfish larvae settling on a coral reef

The fastest swimming fishes in relation to size are found among coral reef fish larvae on their way to settle on reefs. By testing two damselfishes, Chromis atripectoralis and Pomacentrus amboinensis, we show that the high swimming speeds of the pre-settlement larvae are accompanied by the highest rates of oxygen uptake ever recorded in ectothermic vertebrates. As expected, these high rates of oxygen uptake occur at the cost of poor hypoxia tolerance. However, hypoxia tolerance is needed when coral reef fishes seek nocturnal shelter from predators within coral colonies, which can become severely hypoxic microhabitats at night. When the larvae settle on the reef, we found that they go through a striking respiratory transformation, i.e. the capacity for rapid oxygen uptake falls, while the ability for high-affinity oxygen uptake at low oxygen levels is increased. This transition to hypoxia tolerance is needed when they settle on the reef; this was strengthened by our finding that small resident larvae of Acanthochromis polyacanthus, a damselfish lacking a planktonic larval stage, do not display such a transition, being well adapted to hypoxia and showing relatively low maximum rates of oxygen uptake that change little with age.     

A standard unit for monitoring recruitment of fishes to coral reef rubble

We developed a benthic standard monitoring unit for recruitment of fishes (SMURF) to sample fishes that settle in coral rubble and tested it on three fringing reefs on the west coast of Barbados, West Indies. These rubble SMURFs are inexpensive to construct and permit newly settled fishes, including cryptic, secretive and small species, to be quickly and fairly completely (> 83%) removed by divers without returning the unit to the surface. Over a 3-month period involving 9 collections, 32 units on 3 reefs yielded 948 newly settled fishes belonging to 28 taxa (3.3 ± 0.2 S.E. recruits per sample, n = 287) in addition to numerous crustaceans. Most specimens were Sparisoma spp. (Scaridae) (41.8%) and Stegastes partitus (Pomacentridae) (13.6%), with moderate numbers of Scarus spp. (Scaridae) (8.6%), Lythrypnus spp. (Gobiidae) (7.4%), and Pseudogramma gregoryi (Serranidae) (7.2%). Sampling SMURFs at 1-day rather than 11-day intervals yielded approximately twice as many fish, although some taxa were not affected by sampling frequency, indicating taxon-specific differences in post-settlement loss rates. Netting, intended to reduce predation on settlers, did not affect settlement estimates unless algae grew on it, suggesting that the SMURF design minimizes fish predation even in the absence of the netting. Over time, however, the netting resulted in substantial algal growth that was associated with an increase in abundance of Sparisoma spp. and a decrease in the abundance of S. partitus. This suggests that settler estimates are more affected by microhabitat changes from algal growth than by predation. During low settlement periods, SMURFs performed better than light-traps in assessing the daily input of new settlers of two abundant taxa. Eight SMURFs per reef were sufficient to demonstrate differences in settlement rates and temporal correlations in settlement within and among reefs. This study underscores the potential of benthic SMURFs to measure settlement of coral reef fishes at a range of spatio-temporal scales with moderate research cost and effort, and to facilitate the study of the settlement stage of several taxa.