Behavioral and energetic costs of group membership in a coral reef fish

Animals in social aggregations often spend more time foraging than solitary conspecifics. This may be a product of the relative safety afforded by aggregations: group members can devote more time to foraging and less time to antipredator behaviors than solitary animals (the “risk reduction” effect). All else being equal, risk reduction should result in higher food intake for grouped animals. However, intragroup competition may force group members to spend more time foraging in order to obtain the same food ration as solitary individuals (the “resource competition” effect). We compared these opposing explanations of foraging time allocation in a coral reef fish, bluehead wrasse (Thalassoma bifasciatum). Aggregations of juvenile bluehead wrasse experience safety-in-numbers, and preliminary observations suggested that juveniles in aggregations spent more time foraging for copepods in the water column than solitary juveniles. However, the risk reduction and resource competition hypotheses are indistinguishable on the basis of behavioral observations alone. Therefore, we collected behavioral, dietary, and growth data (using otolith growth rings) for bluehead wrasse at multiple reefs around a Caribbean island. Despite spending more time foraging in the water column, grouped fish did not capture more prey items and had slower growth rates than solitary fish. Thus, the increased foraging time of grouped fish appears to reflect resource competition, not risk reduction. This competition may limit the size and frequency of aggregations among juvenile bluehead wrasse, which have been shown to experience reduced mortality rates in larger groups. Bluehead wrasse recruits also spent less time foraging but grew faster at sites where planktonic copepod prey were more abundant. This suggests the possibility that large-scale spatiotemporal variability in the abundance of planktonic copepods over coral reefs may produce corresponding variability in the dynamics of reef fish populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Tracking and mapping sun-synchronous migrations and diel space use patterns of Haemulon sciurus and Lutjanus apodus in the U.S. Virgin Islands

The spatially explicit diel movement patterns of fish using coral reef ecosystems are not well understood, despite the widespread recognition that many common species undergo distinct migrations to utilize different resources during night and day. We used manual acoustic telemetry coupled with global positioning technology to track the detailed spatially explicit daily movements (24 h) of multiple individuals of two common Caribbean fish species, Haemulon sciurus (bluestriped grunt) and Lutjanus apodus (schoolmaster snapper). Movement pathways and day and night activity spaces were mapped and quantified in a Geographic Information System (GIS). Directional sun-synchronous migrations occurred close to astronomical sunset and sunrise. Site fidelity within day and night activity spaces was high. Nine of twelve individuals exhibited overlap of day and night activity spaces and three fish (L. apodus) exhibited complete spatial segregation. Night activity spaces (H. sciurus: 11,309?±?3,548 m²; L. apodus: 9,950?±?3,120 m²) were significantly larger than day activity spaces (H. sciurus: 2,778?±?1,979 m²; L. apodus: 1,291?±?636 m²). The distance between sequential position fixes (step lengths) was significantly greater at night than day, indicative of nocturnal foraging and day resting behavior. Integrating acoustic telemetry, GIS techniques and spatial statistics to study fish movement behavior revealed both individual variability and some broader generality in movement paths and activity spaces suggestive of complex underlying behavioral mechanisms influencing diel movements.     

Multispecies spawning sites for fishes on a low‐latitude coral reef: spatial and temporal patterns

Spawning sites used by one or more species were located by intensively searching nearshore coral reefs of Kimbe Bay (New Britain, Papua New Guinea). Once identified, the spawning sites were surveyed repeatedly within fixed 5 m radius circular areas, for > 2000 h of observations ranging from before dawn to after dusk spanning 190 days between July 2001 and May 2004. A total of 38 spawning sites were identified on the seven study reefs distributed at an average of one site every 60 m of reef edge. Pelagic spawning was observed in 41 fish species from six families. On three intensively studied reefs, all 17 spawning sites identified were used by at least three species, with a maximum of 30 different species observed spawning at a single site. Spawning was observed during every month of the study, on all days of the lunar month, at all states of the tide and at most hours of the day studied. Nevertheless, the majority of species were observed spawning on proportionately more days from December to April, on more days around the new moon and in association with higher tides. The strongest temporal association, however, was with species‐specific diel spawning times spanning < 3 h for most species. While dawn spawning, afternoon spawning and dusk spawning species were differentiated, the time of spawning for the striated surgeonfish Ctenochaetus striatus also differed significantly among sites. The large number of species spawning at the same restricted locations during predictable times suggests that these sites are extremely important on this low‐latitude coral reef.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.     

Using passive acoustic telemetry to infer mortality events in adult herbivorous coral reef fishes

Mortality is considered to be an important factor shaping the structure of coral reef fish communities, but data on the rate and nature of mortality of adult coral reef fishes are sparse. Mortality on coral reefs is intrinsically linked with predation, with most evidence suggesting that predation is highest during crepuscular periods. We tested this hypothesis using passive acoustic telemetry data to determine the time of day of potential mortality events (PMEs) of adult herbivorous reef fishes. A total of 94 fishes were tagged with acoustic transmitters, of which 43 exhibited a PME. Furthermore, we identified five categories of PMEs based on the nature of change in acoustic signal detections from tagged fishes. The majority of PMEs were characterised by an abrupt stop in detections, possibly as a result of a large, mobile predator. Overall, mortality rates were estimated to be approximately 59 % per year using passive acoustic telemetry. The time of day of PMEs suggests that predation was highest during the day and crepuscular periods and lowest at night, offering only partial support for the crepuscular predation hypothesis. Visually oriented, diurnal and crepuscular predators appear to be more important than their nocturnal counterparts in terms of predation on adult reef fishes. By timing PMEs, passive acoustic telemetry may offer an important new tool for investigating the nature of predation on coral reefs.     

Migration of hawksbill turtles Eretmochelys imbricata from Tortuguero, Costa Rica

The hawksbill turtle Eretmochelys imbricata is a widely distributed and critically endangered species that feeds on sponges and fills an important ecological role in the coral reef ecosystem. At Tortuguero, Costa Rica, trend analyses indicate considerable decline in nesting estimated at 77.2–94.5% between 1956 and 2003, as a result of excessive turtle fishing. We analyzed flipper tag returns, satellite telemetry and genetic samples to determine movements and habitat use of adult female Tortuguero hawksbills. Tag returns and satellite telemetry show hawksbills migrate to foraging grounds in Nicaragua and Honduras. Genetic analysis indicates the hawksbills may also migrate to Cuban, Puerto Rican, and possibly Mexican waters. We conclude hawksbills represent an internationally shared resource. There is a close correlation between tag recapture sites, hawksbill foraging grounds and coral reef distribution. Caribbean coral reef decline may reduce food availability and negatively impact hawksbill turtles. Conversely, hawksbill decline may shift the balance on coral reefs by reducing predation pressure on sponges and hence make coral reefs less resilient to natural and anthropogenic threats. Strategies aiming to conserve hawksbills and coral reefs must consider both the extensive hawksbill migrations and the close relationship between the species and the coral reef ecosystem.     

Seasonal occupancy and connectivity amongst nearshore flats and reef habitats by permit Trachinotus falcatus: considerations for fisheries management

We used acoustic telemetry to quantify permit Trachinotus falcatus habitat use and connectivity in proximity to the Florida Keys, USA, and assessed these patterns relative to current habitat and fisheries management practices. From March 2017 to June 2018, 45 permit tagged within 16 km of the lower Florida Keys were detected at stationary acoustic receivers throughout the south Florida region, the majority of which remained within the Special Permit Zone, where more extensive fisheries harvest regulations are implemented. There was a high level of connectivity between nearshore flats (i.e., <3 m water depth) and the Florida reef tract (FRT; 15–40 m water depth), with 75% of individuals detected in both habitats. These locations probably function primarily as foraging and spawning habitats, respectively. Permit occupancy on the FRT peaked during the months of March–September, with the highest number of individuals occurring there in April and May. Specific sites on the FRT were identified as potentially important spawning locations, as they attracted a high proportion of individuals that exhibited frequent visits with high residency durations. There were also significant positive relationships between seasonal habitat-use metrics on the FRT and an empirical permit gonadosomatic index. Large aggregations of permit at spawning sites on the FRT are potentially vulnerable to the effects of fishing (including predation during catch and release) at a critical point in their life cycle. These data on permit space use and movement, coupled with knowledge of stressors on their ecology, provide insights for implementing science-based strategic management plans.     

Non‐reef habitats in a tropical seascape affect density and biomass of fishes on coral reefs

Nonreef habitats such as mangroves, seagrass, and macroalgal beds are important for foraging, spawning, and as nursery habitat for some coral reef fishes. The spatial configuration of nonreef habitats adjacent to coral reefs can therefore have a substantial influence on the distribution and composition of reef fish. We investigate how different habitats in a tropical seascape in the Philippines influence the presence, density, and biomass of coral reef fishes to understand the relative importance of different habitats across various spatial scales. A detailed seascape map generated from satellite imagery was combined with field surveys of fish and benthic habitat on coral reefs. We then compared the relative importance of local reef (within coral reef) and adjacent habitat (habitats in the surrounding seascape) variables for coral reef fishes. Overall, adjacent habitat variables were as important as local reef variables in explaining reef fish density and biomass, despite being fewer in number in final models. For adult and juvenile wrasses (Labridae), and juveniles of some parrotfish taxa (Chlorurus), adjacent habitat was more important in explaining fish density and biomass. Notably, wrasses were positively influenced by the amount of sand and macroalgae in the adjacent seascape. Adjacent habitat metrics with the highest relative importance were sand (positive), macroalgae (positive), and mangrove habitats (negative), and fish responses to these metrics were consistent across fish groups evaluated. The 500‐m spatial scale was selected most often in models for seascape variables. Local coral reef variables with the greatest importance were percent cover of live coral (positive), sand (negative), and macroalgae (mixed). Incorporating spatial metrics that describe the surrounding seascape will capture more holistic patterns of fish–habitat relationships on reefs. This is important in regions where protection of reef fish habitat is an integral part of fisheries management but where protection of nonreef habitats is often overlooked.     

Consumption of coral propagules represents a significant trophic link between corals and reef fish

Mass spawning of corals provides a large seasonal pulse of high-energy prey that potentially benefits reef fish that are capable of capturing and digesting coral propagules. This study examines the range of fish species that consume coral propagules and also tests whether reef fish experience a significant increase in physiological condition when feeding on coral propagules. Thirty-six species of diurnal reef fish were seen to consume coral propagules released during mass coral spawning. Stomach content analyses of three reef fish species (Pomacentrus moluccensis, Abudefduf whitleyi, and Caesio cunning) revealed that both P. moluccensis and A. whitleyi feed almost exclusively on coral propagules during mass coral spawning. Fish feeding extensively on coral propagules also amassed considerable lipid stores, which could greatly improve the quality and survivorship of their progeny. In contrast, C. cunning consumed only very small quantities of coral propagules, and showed no detectable change in lipid stores during the course of the study. This study provides the first direct evidence that reef fish benefit from mass coral spawning, and reveals a potentially significant trophic link between scleractinian corals and reef fish. Accepted: 9 June 2000     

The utility of a long‐term acoustic recording system for detecting white seabass Atractoscion nobilis spawning sounds

This study reports the use of a long‐term acoustic recording system (LARS) to remotely monitor white seabass Atractoscion nobilis spawning sounds at three sites along the southern California coastline, adjacent to Camp Pendleton. On the basis of previous studies of A. nobilis sound production relative to periods of known spawning activity, LARS were set to continuously record ambient sounds for a 2 h period around sunset from April to June 2009. Acoustic analyses identified A. nobilis courtship sounds on 89, 28 and 45% of the days at the three locations, respectively. From 474 h of acoustic data, spawning‐related sounds (chants) were detected on 19 occasions in 2009 with an additional 11 spawning chants recorded during a 2007 validation period. Most spawning chants occurred within 30 min of sunset during the months of May and June at a mean ±s.d . surface temperature of 18·2 ± 1·2° C. Consecutive daily spawning activity was not apparent at any sites in 2009. Atractoscion nobilis spawning chants were recorded at all three sites, suggesting that shallow rocky reefs which support kelp forests provide suitable A. nobilis spawning habitat. Results confirm the utility of passive acoustic recorders for identifying A. nobilis spawning periods and locations.     

Reproductive behavior and mate fidelity in the monogamous goby,Valenciennea longipinnis

Reproductive behavior and mate fidelity of the gobiid fish,Valenciennea longipinnis, were studied on the coral reef at Sesoko Island, Okinawa, Japan. These fish usually live in pairs, not only foraging together for benthic animals in sandy areas, but also constructing several burrows within their home range. Before spawning, both fish, although mainly the male, constructed a mound, piling up dead-coral fragments, pebbles, shells, sand and algae onto one of the burrows. After spawning an egg mass on the ceiling of the burrow, the female stayed outside and continued the construction and maintenance of the mound for 3–5 days until hatching, while the male tended the eggs inside. Mate guarding of females seemed to prevent males from monopolizing several females. Although some pairs showed mate fidelity through several spawnings, more than half of the pairs broke up after only one spawning. The pair bond was broken by mate desertion and the disappearance of each sex. Both sexes preferred larger spawning partners; larger females spawned more eggs and larger males provided better egg care. Mate desertion occurred when larger potential mates, relative to the current partner, became available. The frequency of solitary individuals was higher in males than in females, resulting in females deserting their mates more often than males. Two factors seem to have facilitated mate desertion: (1) occurrence of size mis-matched pairing and (2) overlapping home ranges.     

The application of acoustic Doppler current profilers to measure the timing and patterns of coral larval dispersal

An experiment was conducted along the reefs off west Maui, Hawaii, during the summer of 2003 to monitor the spawning of the reef-building coral Montipora capitata and to determine the role of ocean currents in dispersing the larvae from the natal reef. Instruments documented the environmental forcing during the coral spawning season; drifters were deployed on three successive nights following direct observations of coral spawning. Both the timing and relative magnitude of the coral spawning were identifiable in acoustic backscatter data and correlated to plankton tow data. Each drifter track showed that the surface water containing coral eggs and planula larvae were transported rapidly offshore and not locally retained. Wind and current patterns during the previous year and during subsequent coral spawning events later in the summer were similar to those observed during the drifter releases. This suggests that the trajectories observed during the focused experiment are representative of the general pattern of larval dispersal off west Maui. These findings demonstrate the application of acoustic profilers for remotely imaging coral spawning and predicting their initial dispersal patterns.     

Movement in a large predatory fish: coral trout, Plectropomus leopardus (Pisces: Serranidae), on Heron Reef, Australia

 Movement by the larger more mobile species of coral reef fish plays a significant role in determining patterns in abundance and population structure. Fish movement is also relevant to the use and effectiveness of marine reserves in managing fish populations. Coral trout are large piscivorous serranids supporting major fisheries on the Great Barrier Reef (GBR). This study reports on the within-reef movement of the common coral trout, Plectropomus leopardus, at Heron Reef, southern GBR, over a twelve month period, investigated by tagging and underwater tracking. Tracking of coral trout revealed no apparent relationship between the area moved and stage of tide or time of day. However, movement areas were affected by the size of fish: in spring a linear relationship between fish size and area of movement was measured, but in summer the largest (male) fish moved over significantly smaller areas than medium-sized fish. Movement of males may be related to cleaning behaviour and spawning. Fifty nine percent (n=101) of the tagged fish were resighted over periods of 4–5 months, in “home sites” measuring ∼2000 m². Coral trout were not restricted to home sites, but moved on average 2 km along the reef slope; maximum distances of 7–7.5 km were measured. Coral trout appear to range as mobile, opportunistic predators, but also maintain home sites for access to shelter and cleaning stations. Accepted: 1 August 1996     

Environmental Influences on Patterns of Vertical Movement and Site Fidelity of Grey Reef Sharks (Carcharhinus amblyrhynchos) at Aggregation Sites

We used acoustic telemetry to describe the patterns of vertical movement, site fidelity and residency of grey reef sharks (Carcharhinus amblyrhynchos) on the outer slope of coral reefs in Palau, Micronesia, over a period of two years and nine months. We tagged 39 sharks (mostly adult females) of which 31 were detected regularly throughout the study. Sharks displayed strong inter-annual residency with greater attendance at monitored sites during summer than winter months. More individuals were detected during the day than at night. Mean depths of tagged sharks increased from 35 m in winter to 60 m in spring following an increase in water temperature at 60 m, with maximum mean depths attained when water temperatures at 60 m stabilised around 29°C. Sharks descended to greater depths and used a wider range of depths around the time of the full moon. There were also crepuscular cycles in mean depth, with sharks moving into shallower waters at dawn and dusk each day. We suggest that daily, lunar and seasonal cycles in vertical movement and residency are strategies for optimising both energetic budgets and foraging behaviour. Cyclical patterns of movement in response to environmental variables might affect the susceptibility of reef sharks to fishing, a consideration that should be taken into account in the implementation of conservation strategies.     

Spawning Site Selection and Contingent Behavior in Common Snook,Centropomus undecimalis

Reproductive behavior affects spatial population structure and our ability to manage for sustainability in marine and diadromous fishes. In this study, we used fishery independent capture-based sampling to evaluate where Common Snook occurred in Tampa Bay and if it changed with spawning season, and passive acoustic telemetry to assess fine scale behavior at an inlet spawning site (2007–2009). Snook concentrated in three areas during the spawning season only one of which fell within the expected spawning habitat. Although in lower numbers, they remained in these areas throughout the winter months. Acoustically-tagged snook (n = 31) showed two seasonal patterns at the spawning site: Most fish occurred during the spawning season but several fish displayed more extended residency, supporting the capture-based findings that Common Snook exhibit facultative catadromy. Spawning site selection for iteroparous, multiple-batch spawning fishes occurs at the lifetime, annual, or intra-annual temporal scales. In this study we show colonization of a new spawning site, indicating that lifetime spawning site fidelity of Common Snook is not fixed at this fine spatial scale. However, individuals did exhibit annual and intra-seasonal spawning site fidelity to this new site over the three years studied. The number of fish at the spawning site increased in June and July (peak spawning months) and on new and full lunar phases indicating within population variability in spawning and movement patterns. Intra-seasonal patterns of detection also differed significantly with sex. Common Snook exhibited divergent migration tactics and habitat use at the annual and estuarine scales, with contingents using different overwintering habitat. Migration tactics also varied at the spawning site at the intra-seasonal scale and with sex. These results have important implications for understanding how reproductive behavior affects spatio-temporal patterns of fish abundance and their resilience to disturbance events and fishing pressure.     

Acoustic Telemetry Validates a Citizen Science Approach for Monitoring Sharks on Coral Reefs

Citizen science is promoted as a simple and cost-effective alternative to traditional approaches for the monitoring of populations of marine megafauna. However, the reliability of datasets collected by these initiatives often remains poorly quantified. We compared datasets of shark counts collected by professional dive guides with acoustic telemetry data from tagged sharks collected at the same coral reef sites over a period of five years. There was a strong correlation between the number of grey reef sharks (Carcharhinus amblyrhynchos) observed by dive guides and the telemetry data at both daily and monthly intervals, suggesting that variation in relative abundance of sharks was detectable in datasets collected by dive guides in a similar manner to data derived from telemetry at these time scales. There was no correlation between the number or mean depth of sharks recorded by telemetry and the presence of tourist divers, suggesting that the behaviour of sharks was not affected by the presence of divers during our study. Data recorded by dive guides showed that current strength and temperature were important drivers of the relative abundance of sharks at monitored sites. Our study validates the use of datasets of shark abundance collected by professional dive guides in frequently-visited dive sites in Palau, and supports the participation of experienced recreational divers as contributors to long-term monitoring programs of shark populations.     

Patterns of migration between feeding and spawning sites in a coral reef surgeonfish

Many coral reef fishes exhibit regular localised migrations between feeding and spawning areas, but the factors affecting these migration patterns, such as the distance, frequency and spawning site fidelity are poorly understood. The aim of this study was to investigate the patterns of migration to spawning sites of the surgeonfish, Ctenochaetus striatus (Acanthuridae). We explored relationships amongst an individual’s size and sex, the distance and frequency it migrated from its feeding area to spawning sites, fidelity to particular spawning sites and the number of individuals that aggregated to spawn. In order to achieve this, 406 C. striatus were captured and tagged on inshore reefs in Kimbe Bay (5°30′S 150°6′E), New Britain, Papua New Guinea. Tagged individuals were consistently observed within spatially discrete but overlapping feeding areas (maximum diameter averaging <13 m). The mean distance migrated was 58 m (ranging from 2 to 291 m). No tagged individuals were witnessed spawning at more than one site. Whilst most individuals (n = 88) migrated to the spawning site that was closest to their feeding areas, those that migrated to sites further away (n = 9) always spawned at sites where the number of conspecifics aggregating was larger. Neither the size nor the sex of individuals limited migration distance. However, males migrated significantly more frequently than females (on average once every 2 days vs. once every 3 days), and migration frequency was positively correlated with size in females. Migration distance did not affect the frequency with which individuals spawned. Whether patterns of migration are determined by cost-benefit optimisation, tradition, or an alternative mechanism is unknown.     

Evidence of autumn spawning in Suwannee River Gulf sturgeon,Acipenser oxyrinchus desotoi (Vladykov, 1955)

Evidence of autumn spawning of Gulf sturgeon Acipenser oxyrinchus desotoi in the Suwannee River, Florida, was compiled from multiple investigations between 1986 and 2008. Gulf sturgeon are known from egg collections to spawn in the springtime months following immigration into rivers. Evidence of autumn spawning includes multiple captures of sturgeon in September through early November that were ripe (late‐development ova; motile sperm) or exhibited just‐spawned characteristics, telemetry of fish that made >175 river kilometer upstream excursions to the spawning grounds in September–October, and the capture of a 9.3 cm TL age‐0 Gulf sturgeon on 29 November 2000 (which would have been spawned in late September 2000). Analysis of age‐at‐length data indicates that ca. 20% of the Suwannee River Gulf sturgeon population may be attributable to autumn spawning. However, with the very low sampling effort expended, eggs or early life stages have not yet been captured in the autumn, which would be the conclusive proof of autumn spawning. More sampling, and sampling at previously unknown sites frequented by acoustic telemetry fish, would be required to find eggs.     

Novel species interactions and environmental conditions reduce foraging competency at the temperate range edge of a range-extending coral reef fish

Poleward range extensions of coral reef species can reshuffle temperate communities by generating competitive interactions that did not exist previously. However, novel environmental conditions and locally adapted native temperate species may slow tropical invasions by reducing the ability of invaders to access local resources (e.g. food and shelter). We test this hypothesis on wild marine fish in a climate warming hotspot using a field experiment encompassing artificial prey release. We evaluated seven behaviours associated with foraging and aggressive interactions in a common range-extending coral reef fish (Abudefduf vaigiensis) and a co-shoaling temperate fish (Microcanthus strigatus) along a latitudinal temperature gradient (730 km) in SE Australia. We found that the coral reef fish had reduced foraging performance (i.e. slower prey perception, slower prey inspection, decreased prey intake, increased distance to prey) in their novel temperate range than in their subtropical range. Furthermore, higher abundance of temperate fishes was associated with increased retreat behaviour by coral reef fish (i.e. withdrawal from foraging on released prey), independent of latitude. Where their ranges overlapped, temperate fish showed higher foraging and aggression than coral reef fish. Our findings suggest that lower foraging performance of tropical fish at their leading range edge is driven by the combined effect of environmental factors (e.g. lower seawater temperature and/or unfamiliarity with novel conditions in their extended temperate ranges) and biological factors (e.g. increased abundance and larger body sizes of local temperate fishes). Whilst a future increase in ocean warming is expected to alleviate current foraging limitations in coral reef fishes at leading range edges, under current warming native temperate fishes at their trailing edges appear able to slow the range extension of coral reef fishes into temperate ecosystems by limiting their access to resources.

     

The social organization of adult blue tangs, <Emphasis Type="Italic">Acanthurus coeruleus</Emphasis>, on a fringing reef,Barbados, West Indies

Blue tangs in Barbados exhibit three distinct social modes: territorial, schooling and wandering. We compared the mobility, foraging, aggression performed and received and the use of cleaning stations of adult blue tangs among modes and among habitats within a single fringing reef in Barbados. Evidence from observed switches during focal observations and multiple observations of tagged individuals indicate that fish are either territorial or non-territorial. Non-territorial fish formed schools and wandered. However, wandering can be used during solitary movements by fish in either type. Fish in the territorial mode, not previously described in adults of this species, restricted activity to a small area overlapping the territories of other tangs. They actively chased conspecifics and were chased mostly by damselfishes. They swam more slowly and fed at higher rates than other modes. Fish in the schooling mode ranged widely in compact, polarized groups of conspecifics, congeners and other species. They were not aggressive and were attacked mostly by damselfishes. They swam rapidly and fed at intermediate rates. Fish in the wanderer mode showed neither aggression nor association with other individuals. They swam rapidly, well above the substrate, fed little, were chased by conspecifics, ocean surgeonfish, A. bahianus, and damselfish and visited cleaning stations more often then other modes. All three modes were observed in all four main zones of the reef, and their behaviour changed quantitatively with habitat type. We suggest that territoriality reduces competition for algal food, schooling allows fish to overcome the food defence by damselfish, tangs and ocean surgeonfish, and wandering permits solitary movement over the reef to cleaning stations, feeding sites and other resources.