Long-term effects of the cleaner fish Labroides dimidiatus on coral reef fish communities

Cleaning behaviour is deemed a mutualism, however the benefit of cleaning interactions to client individuals is unknown. Furthermore, mechanisms that may shift fish community structure in the presence of cleaning organisms are unclear. Here we show that on patch reefs (61–285 m²) which had all cleaner wrasse Labroides dimidiatus (Labridae) experimentally removed (1–5 adults reef⁻¹) and which were then maintained cleaner-fish free over 8.5 years, individuals of two site-attached (resident) client damselfishes (Pomacentridae) were smaller compared to those on control reefs. Furthermore, resident fishes were 37% less abundant and 23% less species rich per reef, compared to control reefs. Such changes in site-attached fish may reflect lower fish growth rates and/or survivorship. Additionally, juveniles of visitors (fish likely to move between reefs) were 65% less abundant on removal reefs suggesting cleaners may also affect recruitment. This may, in part, explain the 23% lower abundance and 33% lower species richness of visitor fishes, and 66% lower abundance of visitor herbivores (Acanthuridae) on removal reefs that we also observed. This is the first study to demonstrate a benefit of cleaning behaviour to client individuals, in the form of increased size, and to elucidate potential mechanisms leading to community-wide effects on the fish population. Many of the fish groups affected may also indirectly affect other reef organisms, thus further impacting the reef community. The large-scale effect of the presence of the relatively small and uncommon fish, Labroides dimidiadus, on other fishes is unparalleled on coral reefs.     

Predators reduce abundance and species richness of coral reef fish recruits via non-selective predation

Predators have important effects on coral reef fish populations, but their effects on community structure have only recently been investigated and are not yet well understood. Here, the effect of predation on the diversity and abundance of young coral reef fishes was experimentally examined in Moorea, French Polynesia. Effects of predators were quantified by monitoring recruitment of fishes onto standardized patch reefs in predator-exclosure cages or uncaged reefs. At the end of the 54-day experiment, recruits were 74% less abundant on reefs exposed to predators than on caged ones, and species richness was 42% lower on reefs exposed to predators. Effects of predators varied somewhat among families, however, rarefaction analysis indicated that predators foraged non-selectively among species. These results indicate that predation can alter diversity of reef fish communities by indiscriminately reducing the abundance of fishes soon after settlement, thereby reducing the number of species present on reefs.     

The influence of multiple factors upon reef fish abundance and species richness in a tropical coral complex

The present study was conducted on Tamandaré reefs, northeast Brazil and aimed to analyse the importance of different factors (e.g. tourism activity, fishing activity, coral abundance and algal abundance) on reef fish abundance and species richness. Two distinct reef areas (A ver o mar and Caieiras) with different levels of influence were studied. A total of 8239 reef fish individuals were registered, including 59 species. Site 1 (A ver o mar) presented higher reef fish abundance and richness, with dominance of roving herbivores (29.9 %) and mobile invertebrate feeders (28.7 %). In contrast, at Site 2 (Caieiras) territorial herbivores (40.9 %) predominated, followed by mobile invertebrate feeders (24.6 %). Concerning the benthic community, at Site 1 macroalgae were recorded as the main category (49.3 %); however, Site 2 was dominated by calcareous algae (36.0 %). The most important variable explaining more than 90 % of variance on reef fish abundance and species richness was macroalgae abundance, followed by fishing activity. Phase shifts on coral reefs are evident, resulting in the replacement of coral by macroalgae and greatly influencing reef fish communities. In this context, it is important to understand the burden of the factors that affect reef fish communities and, therefore, influence the extinction vulnerability of coral reef fishes.     

Human impacts on the species-area relationship in reef fish assemblages

The relationship between species richness and area is one of the oldest, most recognized patterns in ecology. Here we provide empirical evidence for strong impacts of fisheries exploitation on the slope of the species–area relationship (SAR). Using comparative field surveys of fish on protected and exploited reefs in three oceans and the Mediterranean Sea, we show that exploitation consistently depresses the slope of the SAR for both power-law and exponential models. The magnitude of change appears to be proportional to fishing intensity. Results are independent of taxonomic resolution and robust across coral and rocky reefs, sampling protocols and statistical methods. Changes in species richness, relative abundance and patch occupancy all appear to contribute to this pattern. We conclude that exploitation pressure impacts the fundamental scaling of biodiversity as well as the species richness and spatial distribution patterns of reef fish. We propose that species–area curves can be sensitive indicators of community-level changes in biodiversity, and may be useful in quantifying the human imprint on reef biodiversity, and potentially elsewhere.     

Ecological structure of assemblages of coral reef fishes on isolated patch reefs

A 9-year study of the structure of assemblages of fish on 20 coral patch reefs, based on 20 non-manipulative censuses, revealed a total of 141 species from 34 families, although 40 species accounted for over 95% of sightings of fish. The average patch reef was 8.5 m² in surface area, and supported 125 fish of 20 species at a census. All reefs showed at least a two-fold variation among censuses in total numbers of fish present, and 12 showed ten-fold variations. There was also substantial variation in the composition and relative abundances of species present on each patch reef, such that censuses of a single patch reef were on average about 50% different from each other in percent similarity of species composition (Czekanowski's index). Species differed substantially in the degree to which their numbers varied from census to census, and in the degree to which their dispersion among patch reefs was modified from census to census. We characterize the 40 most common species with respect to these attributes. The variations in assemblage structure cannot be attributed to responses of fish to a changing physical structure of patch reefs, nor to the comings and goings of numerous rare species. Our results support and extend earlier reports on this study, which have stressed the lack of persistant structure for assemblages on these patch reefs. While reef fishes clearly have microhabitat preferences which are expressed at settlement, the variations in microhabitat offered by the patch reefs are insufficient to segregate many species of fish by patch reef. Instead, at the scale of single patch reefs, and, to a degree, at the larger scale of the 20 patch reefs, most of the 141 species of fish are distributed without regard to differences in habitat structure among reefs, and patterns of distribution change over time. Implications for general understanding of assemblage dynamics for fish over more extensive patches of reef habitat are considered.     

Auditory sensitivity in settlement-stage larvae of coral reef fishes

The larval phase of most species of coral reef fishes is spent away from the reef in the pelagic environment. At the time of settlement, these larvae need to locate a reef, and recent research indicates that sound emanating from reefs may act as a cue to guide them. Here, the auditory abilities of settlement-stage larvae of four species of coral reef fishes (families Pomacentridae, Lutjanidae and Serranidae) and similar-sized individuals of two pelagic species (Carangidae) were tested using an electrophysiological technique, auditory brainstem response (ABR). Five of the six species heard frequencies in the 100–2,000 Hz range, whilst one carangid species did not detect frequencies higher than 800 Hz. The audiograms of the six species were of similar shape, with best hearing at lower frequencies between 100 and 300 Hz. Strong within-species differences were found in hearing sensitivity both among the coral reef species and among the pelagic species. Larvae of the coral reef species had significantly more sensitive hearing than the larvae of the pelagic species. The results suggest that settlement-stage larval reef fishes may be able to detect reef sounds at distances of a few 100 m. If true hearing thresholds are lower than ABR estimates, as indicated in some comparisons of ABR and behavioural methods, the detection distances would be much larger.     

Reef fish community structure of the Fernando de Noronha Archipelago (Equatorial Western Atlantic): the influence of exposure and benthic composition

We studied the reef fish assemblage of eight reefs within the oceanic archipelago of Fernando de Noronha, off northeastern Brazil. In a total of 91 belt transects (20 × 2 m) we recorded 60 species from 28 families. The 25 most abundant species accounted for about 98% of all fish recorded in this study and most of these species are widely distributed in the Western Atlantic. The majority of fish counted were planktivores (37.0%), followed by mobile invertebrate feeders (28.5%), territorial herbivores (11.3%), roving herbivores (10.5%), omnivores (7.1%), macrocarnivores (6.5%) and sessile invertebrate feeders (0.03%). In terms of biomass, roving herbivores were the most representative (41.8%), followed by mobile invertebrate feeders (19.9%), macrocarnivores (14.3%), omnivores (14.0%), piscivores (8.3%), planktivores (1.4%), territorial herbivores (0.3%), and sessile invertebrate feeders (0.03%). Overall, density and biomass of fishes were positively correlated with coral cover and depth, and negatively correlated with wave exposure. These relationships are probably a response to the habitat complexity provided by the higher coral cover in deeper reefs (>10 m) of the archipelago or to the lower water turbulence below 10 m deep. Carnivores and mobile invertebrate feeders were mainly influenced by depth and non-consolidated substratum, planktivores and omnivores by wave exposure and herbivores by algal cover. Although our results suggest that habitat characteristics may play a role in determining the distribution of some fish species, we also found several habitat generalists, suggesting that the community is dominated by versatile species.     

Maintenance of fish diversity on disturbed coral reefs

Habitat perturbations play a major role in shaping community structure; however, the elements of disturbance-related habitat change that affect diversity are not always apparent. This study examined the effects of habitat disturbances on species richness of coral reef fish assemblages using annual surveys of habitat and 210 fish species from 10 reefs on the Great Barrier Reef (GBR). Over a period of 11 years, major disturbances, including localised outbreaks of crown-of-thorns sea star (Acanthaster planci), severe storms or coral bleaching, resulted in coral decline of 46–96% in all the 10 reefs. Despite declines in coral cover, structural complexity of the reef framework was retained on five and species richness of coral reef fishes maintained on nine of the disturbed reefs. Extensive loss of coral resulted in localised declines of highly specialised coral-dependent species, but this loss of diversity was more than compensated for by increases in the number of species that feed on the epilithic algal matrix (EAM). A unimodal relationship between areal coral cover and species richness indicated species richness was greatest at approximately 20% coral cover declining by 3–4 species (6–8% of average richness) at higher and lower coral cover. Results revealed that declines in coral cover on reefs may have limited short-term impact on the diversity of coral reef fishes, though there may be fundamental changes in the community structure of fishes.     

Competition for shelter in a high-diversity system: structure use by large reef fishes

Competition among large reef fishes for shelter beneath tabular structures provides a rare opportunity to study competition in a species-rich environment. The system permits a detailed study of localised competition with major implications for coral reefs with respect to human impacts including climate change. Using underwater video cameras, this study examined competition among 30 species of large reef fishes (from nine families) for access to shelter provided by 26 tabular structures, which may be the highest reported diversity of vertebrates competing for a single resource. Mean concentrations of fishes under tabular structures were also among the highest biomass recorded on reefs (4.71 kg m⁻²). A generated dominance hierarchy for the occupation of shelter appeared to be primarily driven by the size of fishes. In contrast to previous studies, fishes higher in the hierarchy tended to exhibit the lowest levels of aggression. However, size difference between fishes was found to be strongly negatively correlated with the proportion of aggressive interactions (R ² = 0.971, P < 0.0001). The strong competition for the shade provided by these corals highlights concerns about future shifts in the structure of large reef fish communities as corals are lost. This is particularly concerning given the critical functional roles played by certain species of large reef fishes that utilise tabular structure for shelter and which occupy the lower ranks of the dominance hierarchy.

     

Grazing pressure of herbivorous coral reef fishes on low coral-cover reefs

The impact of grazing by herbivorous fishes (Acanthuridae, Scaridae, and Pomacentridae) on low coral-cover reefs was assessed by measuring rates of benthic algal production and consumption on inshore and offshore reefs in the upper Florida Keys. Algal production rates, determined in situ with caged and uncaged experimental plates, were low (mean 1.05 g C m⁻² day⁻¹) and similar among reef types. Algal consumption rates were estimated using two different models, a detailed model incorporating fish bite rates and algal yield-per-bite for one species extrapolated to a guild-wide value, and a general regression relating fish biomass to algal consumption. Algal consumption differed among reef types: a majority of algal production was consumed on offshore reefs (55–100%), whereas consumption on inshore patch reefs was 31–51%. Spatial variation in algal consumption was driven by differences in herbivorous fish species composition, density, and size-structure among reef types. Algal consumption rates also varied temporally due to seasonal declines in bite rates and intermittent presence of large-bodied, vagile, schooling species. Spatial coherence of benthic community structure and temporal stability of algal turf over 3 years suggests that grazing intensity is currently sufficient to limit further spread of macroalgal cover on these low coral-cover reefs, but not to exclude it from the system.     

Determinants of the steep species–area relationship of coral reef fishes

The increase in species richness with area is known as the species–area relationship (SPAR). Although several mutually non-exclusive processes may produce the SPAR, the null, often ignored, hypothesis states that a SPAR can be generated by random placement alone. The log–log-transformed SPAR of coral reef fishes on small patch-reefs revealed a steep slope of 0.55. However, this slope was dependent on the cumulative area of the reef examined and was therefore affected by random placement. After statistically removing the contribution of random placement from the SPAR, the slope was estimated to be 0.21. This is consistent with estimates from other, mostly terrestrial, systems. Furthermore, a randomization procedure, where the probability of fishes to reach a patch was proportional to reef area, showed that the field measured SPAR did not differ from random placement. In addition, fish assemblages on species poor reefs did not form subsets of species rich reefs (i.e., no nestedness) beyond that expected from random placement. Steep log–log-transformed SPARs can be formed by random placement alone, indicating that caution should be used when assigning an ecological meaning to SPARs generated from small spatial scales.     

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.     

Nocturnal relocation of adult and juvenile coral reef fishes in response to reef noise

Juvenile and adult reef fishes often undergo migration, ontogenic habitat shifts, and nocturnal foraging movements. The orientation cues used for these behaviours are largely unknown. In this study, the use of sound as an orientation cue guiding the nocturnal movements of adult and juvenile reef fishes at Lizard Island, Great Barrier Reef was examined. The first experiment compared the movements of fishes to small patch reefs where reef noise was broadcast, with those to silent reefs. No significant responses were found in the 79 adults that were collected, but the 166 juveniles collected showed an increased diversity each morning on the reefs with broadcast noise, and significantly greater numbers of juveniles from three taxa (Apogonidae, Gobiidae and Pinguipedidae) were collected from reefs with broadcast noise. The second experiment compared the movement of adult and juvenile fishes to reefs broadcasting high (>570 Hz), or low (<570 Hz) frequency reef noise, or to silent reefs. Of the 122 adults collected, the highest diversity was seen at the low frequency reefs; and adults from two families (Gobiidae and Blenniidae) preferred these reefs. A similar trend was observed in the 372 juveniles collected, with higher diversity at the reefs with low frequency noises. This preference was seen in the juvenile apogonids; however, juvenile gobiids were attracted to both high and low sound treatments equally, and juvenile stage Acanthuridae preferred the high frequency noises. This evidence that juvenile and adult reef fishes orientate with respect to the soundscape raises important issues for management, conservation and the protection of sound cues used in natural behaviour.     

Fish assemblages on artificial and natural reefs in the Flower Garden Banks National Marine Sanctuary, USA

 Visual censusing was used to characterize fish assemblages on artificial and natural reefs located within the boundaries of the Flower Garden Banks National Marine Sanctuary (FGBNMS) in the northwestern Gulf of Mexico. Emphasis was placed on determining spatial and temporal patterns in habitat utilization by fishes on an offshore artificial reef (Mobil Platform HI-A389A). Overall, 43 species were observed during diurnal surveys in the upper 24 m of the artificial reef. Midwater pelagic fishes (i.e., carangids and scombrids) accounted for over 50% of all taxa enumerated on the artificial reef; however, these taxa were transient members of the assemblage and were observed infrequently. Labrids, pomacentrids, and serranids were the dominant reef-dependent taxa. Distinct trends in vertical, diel, and seasonal abundances were observed for juvenile and adult fishes. Of the three designated depth zones (upper 1.5–9.0, middle 9.0–16.5; lower 16.5–24.0 m), abundance and species diversity were lowest in the upper zone. Nocturnal counts were characterized by a marked reduction or complete absence of most species, due in part to twilight cover-seeking and movement activities. Seasonal variation in community composition and species abundance (May versus September) was primarily due to recruitment of juveniles (0-age fishes) to the artificial reef in late summer. Increases in total fish abundance (all taxa combined) coincided with both increasing habitat rugosity and degree of fouling. Species richness on natural coral reefs in the FGBNMS was higher than on the artificial reef. Unlike the artificial reef, fish assemblages on the natural reefs were dominated by a single family (Pomacentridae) which accounted for over 50% of all individuals observed. Accepted: 1 August 1996     

Identifying the ichthyoplankton of a coral reef using DNA barcodes

Marine fishes exhibit spectacular phenotypic changes during their ontogeny, and the identification of their early stages is challenging due to the paucity of diagnostic morphological characters at the species level. Meanwhile, the importance of early life stages in dispersal and connectivity has recently experienced an increasing interest in conservation programmes for coral reef fishes. This study aims at assessing the effectiveness of DNA barcoding for the automated identification of coral reef fish larvae through large‐scale ecosystemic sampling. Fish larvae were mainly collected using bongo nets and light traps around Moorea between September 2008 and August 2010 in 10 sites distributed in open waters. Fish larvae ranged from 2 to 100 mm of total length, with the most abundant individuals being <5 mm. Among the 505 individuals DNA barcoded, 373 larvae (i.e. 75%) were identified to the species level. A total of 106 species were detected, among which 11 corresponded to pelagic and bathypelagic species, while 95 corresponded to species observed at the adult stage on neighbouring reefs. This study highlights the benefits and pitfalls of using standardized molecular systems for species identification and illustrates the new possibilities enabled by DNA barcoding for future work on coral reef fish larval ecology.     

Species composition,habitat configuration and seasonal changes of coral reef fish assemblages in western Mexico

In spite of their ecological and economic importance, reef fishes from the coast of Oaxaca, Mexico are rarely studied, therefore precluding their management and conservation. In order to identify the set of habitat characteristics/environmental conditions that predict major shifts in fish assemblages in space and time, a stationary census (5′, φ = 5 m) was conducted on a semi‐monthly basis from 2006 to 2009 at patch reefs along the coast. Habitat configuration was gathered using 25 m long point‐intersect transects (data every 25 cm), recording all underlying coral species and substrate characteristics (rocks, sand, algal mats, rubble or dead corals). Recorded were 65 452 fishes grouped in 11 orders, 36 families, 65 genera and 89 species. Labridae (nine species), Pomacentridae (eight species) and Serranidae (seven species) were the most frequent families. Abundance is severely skewed among species; four species Thalassoma lucasanum, Chromis atrilobata, Apogon pacificus and Stegastes acapulcoensis comprise nearly 59% of the fish abundance, 11 species contribute 30%, whereas most of the species (75) can be considered as rare since they contribute <1% each to the total. Species richness and family‐level assemblage composition are similar to those recorded elsewhere in the eastern Pacific. Non‐parametric multivariate analysis of variance demonstrated that changes of diversity metrics might be associated with environmental differences on the scale of hundreds of meters to kilometers, as well as coupled with major changes on oceanographic variables throughout time, exerting meaningful changes on reef‐related fish assemblages.     

Relationships Between Reef Fish Communities and Remotely Sensed Rugosity Measurements in Biscayne National Park,Florida, USA

The realization that coral reef ecosystem management must occur across multiple spatial scales and habitat types has led scientists and resource managers to seek variables that are easily measured over large areas and correlate well with reef resources. Here we investigate the utility of new technology in airborne laser surveying (NASA Experimental Advanced Airborne Research Lidar (EAARL)) in assessing topographical complexity (rugosity) to predict reef fish community structure on shallow (<10 m deep) patch reefs. Marine portions of Biscayne National Park, Florida, USA, were surveyed remotely using the EAARL, and reef fish populations were visually surveyed on 10 patch reefs at independent, randomly selected stations (n = 10–13 per reef). Rugosity at each station was assessed in situ by divers using the traditional chain-transect method (10-m scale), and remotely using the EAARL submarine topography data at multiple spatial scales (2, 5, and 10 m). The rugosity and biological datasets were analyzed together to elucidate the predictive power of EAARL rugosity in describing the variance in reef fish community variables and to assess the correlation between chain-transect and EAARL rugosity. EAARL rugosity was not well correlated with chain-transect rugosity, or with species richness of fishes (although statistically significant, the amount of variance explained by the model was very low). Variance in reef fish community attributes was better explained in reef-by-reef variability than by physical variables. However, once the reef-by-reef variability was taken into account in a two-way analysis of variance, the importance of rugosity could be seen on individual reefs. Fish species richness and abundance were statistically higher at high rugosity stations compared to medium and low rugosity stations, as predicted by prior ecological research. The EAARL shows promise as an important mapping tool for reef resource managers as they strive to inventory and protect coral reef resources.     

Counts of coral reef fishes by an experienced observer are not biased by the number of target species

Fishes are commercially, recreationally and functionally important inhabitants of coral reefs. Accordingly, accurate assessments of fish abundance and diversity are necessary for effective reef management. While some reef fish monitoring programmes target all fishes, many survey a subset of common and visually obvious species. Changing to counts of all species, while desirable, risks new and unwelcome biases from altered observer swimming speeds and search patterns. Here we test whether substantially increasing the number of target species in an established fish monitoring programme biases counts of the original subset, so precluding ongoing comparisons with historical data. A subset of 141 fish species have been visually surveyed along 50 × 5 m transects over 27 years throughout Australia's Great Barrier Reef. We experimentally compared counts of the subset from standard subset-only surveys and from surveys of all species (excluding small site-attached fishes that are surveyed separately) at three diverse reefs. Subset species richness and abundance, in total and of major families, and assemblage structure did not differ due to survey method. The high-level experience of the one observer appeared to overcome new biases from counting more species and the extra 2 min/transect was not logistically excessive. Surveys of all fishes recorded almost 80% more species than subset-only surveys and >130% higher total abundance on average from >150% more genera that included abundant and functionally important taxa. Overall, fish counts by an experienced observer were not biased by the number of species surveyed and counts of all species markedly improved assessments of reef fish diversity and function.     

Juvenile coral reef fish use sound to locate habitats

There is limited knowledge of the orientation cues used by reef fish in their movement among different habitats, especially those cues used during darkness. Although acoustic cues have been found to be important for settlement-stage fish as they seek settlement habitats, only a small number of studies support the possible role of acoustic cues in the orientation of post-settled and adult reef fish. Therefore, the aim of this study was to determine whether habitat-specific acoustic cues were involved in the nocturnal movements of juvenile reef fish to small experimental patch reefs that were broadcasting sound previously recorded from different habitats (Fringing Reef, Lagoon, Silent). Juvenile fish arriving at each patch reef were caught the next morning by divers and were identified. There were a greater number of occasions when juvenile fish (from all species together) moved onto the patch reefs broadcasting Fringing Reef and Lagoon sound (43 and 38%, respectively) compared to Silent reefs (19%) (χ² = 33.5; P < 0.05). There were significantly more occasions when juvenile fish from the family Nemipteridae were attracted to the patch reefs broadcasting Lagoon sound (63%) versus those reefs broadcasting either Fringing Reef sound (31%) or Silent (6%). In contrast, there were more occasions when juveniles from the family Pomacentridae were attracted to the patch reefs broadcasting Fringing Reef sound (56%) than either Lagoon (24%) or Silent patch reefs (20%) (χ² = 19.5; P < 0.05). These results indicate that some juvenile fish use specific habitat sounds to guide their nocturnal movements. Therefore, the fish are able to not only use the directional information contained in acoustic cues, but can also interpret the content of the acoustic signals for relevant habitat information which is then used in their decision-making for orientation.     

Near-surface enrichment of zooplankton over a shallow back reef: implications for coral reef food webs

Zooplankton were 3–8 times more abundant during the day near the surface than elsewhere in the water column over a 1–2.4 m deep back reef in Moorea, French Polynesia. Zooplankton were also significantly more abundant near the surface at night although gradients were most pronounced under moonlight. Zooplankton in a unidirectional current became concentrated near the surface within 2 m of departing a well-mixed trough immediately behind the reef crest, indicating that upward swimming behavior, rather than near-bottom depletion by reef planktivores, was the proximal cause of these gradients. Zooplankton were highly enriched near the surface before and after a full lunar eclipse but distributed evenly throughout the water column during the eclipse itself supporting light as a proximal cue for the upward swimming behavior of many taxa. This is the first investigation of the vertical distribution of zooplankton over a shallow back reef typical of island barrier reef systems common around the world. Previous studies on deeper fringing reefs found zooplankton depletion near the bottom but no enrichment aloft. In Moorea, where seawater is continuously recirculated out the lagoon and back across the reef crest onto the back reef, selection for upward swimming behavior may be especially strong, because the surface serves both as a refuge from predation and an optimum location for retention within the reef system. Planktivorous fish and corals that can forage or grow even marginally higher in the water column might have a substantial competitive advantage over those nearer the bottom on shallow reefs. Zooplankton abundance varied more over a few tens of centimeters vertical distance than it did between seasons or even between day and night indicating that great care must be taken to accurately assess the availability of zooplankton as food on shallow reefs.