Wave disturbance overwhelms top-down and bottom-up control of primary production in California kelp forests

We took advantage of regional differences in environmental forcing and consumer abundance to examine the relative importance of nutrient availability (bottom-up), grazing pressure (top-down), and storm waves (disturbance) in determining the standing biomass and net primary production (NPP) of the giant kelp Macrocystis pyrifera in central and southern California. Using a nine-year data set collected from 17 sites we show that, despite high densities of sea urchin grazers and prolonged periods of low nutrient availability in southern California, NPP by giant kelp was twice that of central California where nutrient concentrations were consistently high and sea urchins were nearly absent due to predation by sea otters. Waves associated with winter storms were consistently higher in central California, and the loss of kelp biomass to winter wave disturbance was on average twice that of southern California. These observations suggest that the more intense wave disturbance in central California limited NPP by giant kelp under otherwise favorable conditions. Regional patterns of interannual variation in NPP were similar to those of wave disturbance in that year-to-year variation in disturbance and NPP were both greater in southern California. Our findings provide strong evidence that regional differences in wave disturbance overwhelmed those of nutrient supply and grazing intensity to determine NPP by giant kelp. The important role of disturbance in controlling NPP revealed by our study is likely not unique to giant kelp forests, as vegetation dynamics in many systems are dominated by post-disturbance succession with climax communities being relatively uncommon. The effects of disturbance frequency may be easier to detect in giant kelp because it is fast growing and relatively short lived, with cycles of disturbance and recovery occurring on time scales of years. Much longer data sets (decades to centuries) will likely be needed to properly evaluate the role of disturbance relative to other processes in determining patterns of NPP in other systems.     

Trophic cascades result in large-scale coralline algae loss through differential grazer effects

Removal of predators can have strong indirect effects on primary producers through trophic cascades. Crustose coralline algae (CCA) are major primary producers worldwide that may be influenced by predator removal through changes in grazer composition and biomass. CCA have been most widely studied in Caribbean and temperate reefs, where cover increases with increasing grazer biomass due to removal of competitive fleshy algae. However, each of these systems has one dominant grazer type, herbivorous fishes or sea urchins, which may not be functionally equivalent. Where fishes and sea urchins co-occur, fishing can result in a phase shift in the grazing community with subsequent effects on CCA and other substrata. Kenyan reefs have herbivorous fishes and sea urchins, providing an opportunity to determine the relative impacts of each grazer type and evaluate potential human-induced trophic cascades. We hypothesized that fish benefit CCA, abundant sea urchins erode CCA, and that fishing indirectly reduces CCA cover by removing sea urchin predators. We used closures and fished reefs as a large-scale, long-term natural experiment to assess how fishing and resultant changes in communities affect CCA abundance. We used a short-term caging experiment to directly test the effects of grazing on CCA. CCA cover declined with increasing fish and sea urchin abundance, but the negative impact of sea urchin grazing was much stronger than that of fishes. Abundant sea urchins reduced the CCA growth rate to almost zero and prevented CCA accumulation. A warming event (El Ni?o Southern Oscillation, ENSO) occurred during the 18-year study and had a strong but short-term positive effect on CCA cover. However, the effect of the ENSO on CCA was lower in magnitude than the effect of sea urchin grazing. We compare our results with worldwide literature on bioerosion by fishes and sea urchins. Grazer influence depends on whether benefits of fleshy algae removal outweigh costs of grazer-induced bioerosion. However, the cost-benefit ratio for CCA appears to change with grazer type, grazer abundance, and environment. In Kenya, predator removal leads to a trophic cascade that is expected to reduce net calcification of reefs and therefore reduce reef stability, growth, and resilience.     

Mass mortality ofDiadema antillarum

It has been hypothesized that herbivorous fishes and the regular echinoidDiadema antillarum Philippi compete for benthic algae as their major food resource. Mass mortality ofD. antillarum in February 1984 provided the opportunity to test the hypothesis that herbivorous fishes and sea urchins were competing previously. Visual censuses of herbivorous fishes conducted over 4 yr in four reef zones on Tague Bay Reef, St. Croix, U.S. Virgin Islands, before and after the mass mortality indicated that population densities increased approximately three-fold in backreef and shallow (2m) forereef zones and two-fold, and four-fold in mid (5m) and deep (10m) forereef zones, respectively. Juvenile parrotfishes constituted the major component of these increases, except in the shallow forereef where acanthurids became most abundant. Grazing intensity by herbivorous fishes increased in three of the four reef zones immediately following the mass mortality. These data support the hypothesis that exploitative competition for algal resources was occurring prior to the sea urchin mass-mortality, although alternative hypotheses cannot be discounted completely. Despite the increases in the abundances of, and grazing by, herbivorous fishes, the algal community continued to increase in percent cover and biomass, indicating that increased grazing by fishes does not compensate for the loss of grazing byD. antillarum in controlling algal abundance and community structure.     

Territorial User Rights for Fisheries as Ancillary Instruments for Marine Coastal Conservation in Chile

Territorial user rights for fisheries have been advocated as a way to achieve sustainable resource management. However, few researchers have empirically assessed their potential as ancillary marine conservation instruments by comparing them to no‐take marine protected areas. In kelp (Lessonia trabeculata) forests of central Chile, we compared species richness, density, and biomass of macroinvertebrates and reef fishes among territorial‐user‐right areas with low‐level and high‐level enforcement, no‐take marine protected areas, and open‐access areas in 42 100‐m subtidal transects. We also assessed structural complexity of the kelp forest and substratum composition. Multivariate randomized permutation tests indicated macroinvertebrate and reef fish communities associated with the different access regimes differed significantly. Substratum composition and structural complexity of kelp forest did not differ among access regimes. Univariate analyses showed species richness, biomass, and density of macroinvertebrates and reef fishes were greater in highly enforced territorial‐user‐right areas and no‐take marine protected areas than in open‐access areas. Densities of macroinvertebrates and reef fishes of economic importance were not significantly different between highly enforced territorial‐user‐right and no‐take marine protected areas. Densities of economically important macroinvertebrates in areas with low‐level enforcement were significantly lower than those in areas with high‐level enforcement and no‐take marine protected areas but were significantly higher than in areas with open access. Territorial‐user‐right areas could be important ancillary conservation instruments if they are well enforced. Derechos de Usuario Territoriales para Pesquerías como Instrumentos Accesorios para la Conservación Marina Costera en Chile     

Gizzard shad put a freeze on winter mortality of age-0 yellow perch but not white perch.

Four decades of observations on the limnology and fishes of Oneida Lake, New York, USA, provided an opportunity to investigate causes of mortality during winter, a period of resource scarcity for most juvenile fishes, in age-0 yellow perch (Perca flavescens) and age-0 white perch (Morone americana). This time series contains several environmental (e.g., winter severity) and biological (e.g., predator abundance) signals that can be used to disentangle multiple effects on overwinter mortality of these fishes. A multiple regression analysis indicated that age-0 yellow perch winter mortality was inversely related to fish length in autumn and to the abundance of alternative prey (gizzard shad [Dorosoma cepedianum] and white perch). However, no length-selective predation of yellow perch by one of the main predators, adult walleye (Sander vitreus), was detected. In contrast, white perch mortality was directly associated with total predator biomass and abundance of white perch in autumn, and inversely related to yellow perch abundance as a potential buffer species, but not to the abundance of gizzard shad. Winter severity was not a significant predictor of mortality for either perch species. Predicted winter starvation mortality, from a model described in the literature, was much lower than observed mortality for yellow perch. Similar models for white perch were correlated with observed mortality. These results collectively suggest that predation is the main mechanism shaping winter mortality of yellow perch, while both predation and starvation may be important for white perch. This analysis also revealed that gizzard shad buffer winter mortality of yellow perch. Although winter duration determines the northern limit of fish distributions, in mid-latitude Oneida Lake and for these species, predator-prey interactions seem to exert a greater influence on winter mortality than starvation.     

Exploited species impacts on trophic linkages along reef-seagrass interfaces in the Florida Keys

The removal of fish biomass by extensive commercial and recreational fishing has been hypothesized to drastically alter the strength of trophic linkages among adjacent habitats. We evaluated the effects of removing predatory fishes on trophic transfers between coral reefs and adjacent seagrass meadows by comparing fish community structure, grazing intensity, and invertebrate predation potential in predator-rich no-take sites and nearby predator-poor fished sites in the Florida Keys (USA). Exploited fishes were more abundant at the no-take sites than at the fished sites. Most of the exploited fishes were either omnivores or invertivores. More piscivores were recorded at no-take sites, but most (approximately 95%) were moderately fished and unexploited species (barracuda and bar jacks, respectively). Impacts of these consumers on lower trophic levels were modest. Herbivorous and smaller prey fish (< 10 cm total length) densities and seagrass grazing diminished with distance from reefs and were not negatively impacted by the elevated densities of exploited fishes at no-take sites. Predation by reef fishes on most tethered invertebrates was high, but exploited species impacts varied with prey type. The results of the study show that, even though abundances of reef-associated fishes have been reduced at fished sites, there is little evidence that this has produced cascading trophic effects or interrupted cross-habitat energy exchanges between coral reefs and seagrasses.     

Regulation of algal community development in a Macrocystis pyrifera forest

The effects of small and large-scale roughness, overstory development, competition for space with sessile animals, and grazing on algal community development in a subtidal Macrocystis pyrifera forest were examined using specially prepared concrete blocks as substrata. Variation in small-scale roughness (crevices and grooves in the order of 0.1 to 3 mm width and depth) had no significant effects on community composition. However, M. pyrifera colonization, algal diversity, and sessile animal biomass were higher near the upper horizontal edges of blocks and concrete prisms. This “edge” effect may result from a combination of increased spore and larval settlement and enhanced growth of plants and animals associated with the turbulent eddies formed around these obstructions. Natural and experimentally produced variations in the algal overstory demonstrated that the presence of an overstory can reduce algal diversity and cover beneath. Caging experiments suggest that predatory fishes and sea-stars indirectly affect the algal community by removing sessile animals (primarily bryozoans) which compete with the algae for space. The exclusion of grazers resulted in increased growth of Gigartina spp. Selective grazing on this genus may account for its reduced abundance in the study area.     

Fishing‐gear restrictions and biomass gains for coral reef fishes in marine protected areas

Considerable empirical evidence supports recovery of reef fish populations with fishery closures. In countries where full exclusion of people from fishing may be perceived as inequitable, fishing‐gear restrictions on nonselective and destructive gears may offer socially relevant management alternatives to build recovery of fish biomass. Even so, few researchers have statistically compared the responses of tropical reef fisheries to alternative management strategies. We tested for the effects of fishery closures and fishing gear restrictions on tropical reef fish biomass at the community and family level. We conducted 1,396 underwater surveys at 617 unique sites across a spatial hierarchy within 22 global marine ecoregions that represented 5 realms. We compared total biomass across local fish assemblages and among 20 families of reef fishes inside marine protected areas (MPAs) with different fishing restrictions: no‐take, hook‐and‐line fishing only, several fishing gears allowed, and sites open to all fishing gears. We included a further category representing remote sites, where fishing pressure is low. As expected, full fishery closures, (i.e., no‐take zones) most benefited community‐ and family‐level fish biomass in comparison with restrictions on fishing gears and openly fished sites. Although biomass responses to fishery closures were highly variable across families, some fishery targets (e.g., Carcharhinidae and Lutjanidae) responded positively to multiple restrictions on fishing gears (i.e., where gears other than hook and line were not permitted). Remoteness also positively affected the response of community‐level fish biomass and many fish families. Our findings provide strong support for the role of fishing restrictions in building recovery of fish biomass and indicate important interactions among fishing‐gear types that affect biomass of a diverse set of reef fish families.     

Effects of kinship on growth of a live-bearing reef fish

The influence of genetic relatedness on the individual performance (e.g. growth, development) of animals is often tied to agonistic or cooperative behaviors among conspecifics, and studies of the effects of kinship have produced mixed results. To explore genetic relatedness independent of these behaviors, we investigated the effects of kinship on the growth of the kelp perch Brachyistius frenatus, a live-bearing, planktivorous marine reef fish that is capable of only limited dispersal. Although juveniles occur in aggregations and compete for food resources, they do not exhibit overt aggressive or cooperative behavioral interactions. We hypothesized that under competition and in the absence of these behaviors, sibling and non-sibling groups of juvenile B. frenatus raised at the same densities in the field would not differ in average growth, but that siblings would exhibit lower variation in growth, simply due to genetic similarities in inherent growth rates. Pregnant, female kelp perch were collected and placed in cages until parturition was complete. Groups of young, recently born from the same mother or from different mothers, were then raised in the field for 9 wk. Our results revealed that average growth rates were similar between sibling and non-sibling treatments. While variation in growth increased initially in non-siblings, siblings showed little such variation. This divergence, however, was not consistent over the duration of the experiment, and variation in the growth of siblings ultimately converged with that of non-siblings. Effects of genetic relatedness would be most likely to manifest themselves early after birth, before environmental factors exert their influence, and this may explain the initial separation but eventual convergence in variation in growth between sibling and non-sibling treatments. For B. frenatus and other organisms that will encounter relatives and compete for resources without overt behavioral interactions, the degree of kinship may play a minor role in the demography of local populations. Received: 26 January 1998 / Accepted: 30 September 1998     

Direct versus indirect methods of quantifying herbivore grazing impact on a coral reef

Two methods were used to assess the grazing impact of roving herbivorous fishes across a coral reef depth gradient within Pioneer Bay, Orpheus Island, Great Barrier Reef. The first technique employed was a method traditionally used to quantify herbivory on coral reefs via the (indirect) inference of herbivore impact from biomass estimates and reported feeding rates. The second method (one of a range of direct approaches) used remote underwater video cameras to film the daily feeding activity of roving herbivores in the absence of divers. Both techniques recorded similar patterns and relative levels of herbivore biomass across five reef zones at the study site. Indirect estimates of the grazing impact across the reef depth gradient of the three predominant species of herbivore broadly coincided with levels quantified directly by remote underwater video, indicating that, to a large extent, presence does correspond to function. However, the video data suggested that, for individual species in particular reef zones, the absolute level of impact may be less than that inferred from presence. In the case of the parrotfish Scarus rivulatus, the video recordings suggested that, at the reef crest, an average of 52% (±18 SE) of each m² area of reef would be grazed each month, compared with an area of 109% (±41 SE) suggested by inferring grazing activity from presence alone. Potential biases associated with remote video recorders may explain some of the discrepancy between values. Overall, the results suggest that, for some fish groups, the indirect method of inferring function from presence can provide a good indication of relative levels of herbivore impact across a coral reef. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of Human Population Density and Proximity to Markets on Coral Reef Fishes Vulnerable to Extinction by Fishing

Coral reef fisheries are crucial to the livelihoods of tens of millions of people; yet, widespread habitat degradation and unsustainable fishing are causing severe depletion of stocks of reef fish. Understanding how social and economic factors, such as human population density, access to external markets, and modernization interact with fishing and habitat degradation to affect fish stocks is vital to sustainable management of coral reef fisheries. We used fish survey data, national social and economic data, and path analyses to assess whether these factors explain variation in biomass of coral reef fishes among 25 sites in Solomon Islands. We categorized fishes into 3 groups on the basis of life‐history characteristics associated with vulnerability to extinction by fishing (high, medium, and low vulnerability). The biomass of fish with low vulnerability was positively related to habitat condition. The biomass of fishes with high vulnerability was negatively related to fishing conducted with efficient gear. Use of efficient gear, in turn, was strongly and positively related to both population density and market proximity. This result suggests local population pressure and external markets have additive negative effects on vulnerable reef fish. Biomass of the fish of medium vulnerability was not explained by fishing intensity or habitat condition, which suggests these species may be relatively resilient to both habitat degradation and fishing. Efectos de la Densidad de Poblaciones Humanas y la Proximidad del Mercado sobre Peces de Arrecifes de Coral Vulnerables a la Extinción     

Herbivory by the Caribbean king crab on coral patch reefs

Caribbean coral reefs are increasingly dominated by macroalgae instead of corals due to several factors, including the decline of herbivores. Yet, virtually unknown is the role of crustacean macrograzers on coral reef macroalgae. We examined the effect of grazing by the Caribbean king crab (Mithrax spinosissimus) on coral patch reef algal communities in the Florida Keys, Florida (USA), by: (1) measuring crab selectivity and consumption of macroalgae, (2) estimating crab density, and (3) comparing the effect of crab herbivory to that of fishes. Mithrax prefers fleshy macroalgae, but it also consumes relatively unpalatable calcareous algae. Per capita grazing rates by Mithrax exceed those of most herbivorous fish, but Mithrax often occurs at low densities on reefs and its foraging activities are reduced in predator-rich environments. Therefore, the effects of grazing by Mithrax tend to be localized and when at low density contribute primarily to spatial heterogeneity in coral reef macroalgal communities.     

Behavioral response to predators reverses the outcome of competition between prey species

Summary In a laboratory experiment it was shown that piscivorous predators reversed the outcome of competitive interactions between two fish prey species, juveniles of roach (Rutilus rutilus) and perch (Perca fluviatilis), by behaviorally affecting their use of two available habitats, an open water habitat and a structurally complex refuge. The shift in the competitive relationship was the result of predators forcing the juvenile fishes into a prey refuge with high structural complexity. While roach was competitively superior in the unstructured habitat, perch was superior in the structurally complex prey refuge. The reversal in competitive relationship was demonstrated both with respect to foraging rate and growth rate and resulted from the high structural complexity in the prey refuge interfering with the roach's swimming performance. Because survival and growth patterns through the juvenile stages have profound effects on the population/community dynamics of size-structured populations such as those of fish, behaviorally induced changes in competitive ability should have significant implications also at the population and community levels.     

Simulating the potential for ecological restoration of dryland forests in Mexico under different disturbance regimes

Examining the potential for ecological restoration is important in areas where anthropogenic disturbance has degraded forest landscapes. However, the conditions under which restoration of degraded tropical dry forests (TDF) might be achieved in practice have not been determined in detail. In this study, we used LANDIS-II, a spatially explicit model of forest dynamics, to assess the potential for passive restoration of TDF through natural regeneration. The model was applied to two Mexican landscapes under six different disturbance regimes, focusing on the impact of fire and cattle grazing on forest cover, structure and composition. Model results identified two main findings. First, tropical dry forests are more resilient to anthropogenic disturbance than expected. Results suggested that under both a scenario of small, infrequent fires and a scenario of large, frequent fires, forest area can increase relatively rapidly. However, forest structure and composition differed markedly between these scenarios. After 400 years, the landscape becomes increasingly occupied by relatively shade-tolerant species under small, infrequent fires, but only species with both relatively high shade tolerance and high fire tolerance can thrive under conditions with large, frequent fires. Second, we demonstrated that different forms of disturbance can interact in unexpected ways. Our projections revealed that when grazing acts in combination with fire, forest cover, structure and composition vary dramatically depending on the frequency and extent of the fires. Results indicated that grazing and fire have a synergistic effect causing a reduction in forest cover greater than the sum of their individual effects. This suggests that passive landscape-scale restoration of TDF is achievable in both Mexican study areas only if grazing is reduced, and fires are carefully managed to reduce their frequency and intensity.     

Effects of algal grazing and aggressive behaviour of the fishes Pomacentrus lividus and Acanthurus sohal on coral-reef ecology

Aggressive behaviour of the fishes Pomacentrus lividus Bl. Schn. and Acanthurus sohal Forskal from the Red Sea is briefly described, and its effect on intensity of algal grazing by herbivorous fish is demonstrated by settlement experiments. Green filamentous alga settles and grows at shallow depths over large areas of coral reefs, but is cropped by fishes to such an extent that it forms only a thin patchy matting on dead corals. Within pomacentrid territories, the alga forms a thicker matting on loosely cemented coralline rubble. Optimum depth range for growth occurs at less than 20 m. Rich growths of green filamentous alga, such as those which occur within pomacentrid territories or on settlement plates protected by wire netting cages, inhibit settlement of “lithothamnion” and invertebrates. While rasping and grazing fish feeders such as parrot fish and surgeon fish limit the distribution of certain invertebrates such as spirorbids, in shallow water it is also true that, were it not for such active removal of green filamentous alga, “lithothamnion” and many invertebrates would find ewer surfaces suitable for settlement.     

The effect of predation on artificial reef juvenile demersal fish species

There is a concern that artificial reefs (AR) may act purely as fishing aggregation devices. Predators attracted to ARs can influence the distribution and abundance of prey fish species. Determining the role of predators in AR is important in advancing the understanding of community interactions. This paper documents the effects of predation on fish assemblages of AR located near a coastal lagoon fish nursery. The Dicentrarchus labrax is a very opportunistic species preying on juveniles (0⁺ and 1⁺ age classes) of several demersal fish species on the ARs. Reef prey and sea bass abundance were negatively correlated. The mean numbers of prey per sea bass stomach increased with the increase of reef fish prey abundance, suggesting that predation has a significant influence, resulting in a decrease in prey abundance. Prey mortality (4–48%) of demersal reef fish associated species depends on bass density. Prey selection was related both with prey abundance and vulnerability. Results showed that D. labrax predation on AR-fish associated species can increase prey natural mortality. However, the role of bass predation on the ecological functioning of exploited ARs is not clear. There may be increases in local fishing yields due either to an increase in predator biomass through aggregation of sea bass attracted to ARs or to greater production. In contrast, predation on juveniles of economically important reef fish preys, especially the most frequent and abundant (Boops boops), can contribute to a decrease in recruitment to the fishery. Our results indicate that inter-specific interactions (predator–prey) are important in terms of conservation and management, as well as for the evaluation of the long-term effects of reef deployment. Thus, it is necessary to consider ecological interactions, such as predation, prior to the development and deployment of artificial habitats as a tool for rehabilitation.     

Scale-dependent changes in the importance of larval supply and habitat to abundance of a reef fish

While there is great interest in the degree to which local interactions "scale-up" to predict regional patterns of abundance, few studies in marine systems have simultaneously examined patterns of abundance at both the large scale (tens of kilometers) typical of larval movement and the small scale (meters) typical of post-settlement interactions. We addressed this gap by monitoring larval supply, adult survivorship, and giant kelp (Macrocystis pyrifera, a primary habitat-forming species) abundance for 13 populations of kelp bass (Paralabrax clathratus) spread over approximately 200 km in the Santa Barbara Channel, California, USA. At the small, within-site scale, both recruitment and adult survivorship of kelp bass were density-dependent and positively related to kelp abundance. At the larger, among-site scale, the spatial pattern of adult kelp bass abundance was predicted well by the pattern of kelp bass larval supply, but there was a consistent negative spatial relationship between kelp abundance and kelp bass larval supply despite the positive effects of kelp on kelp bass at the smaller spatial scale. This large-scale negative relationship was likely a product of a channel-wide spatial mismatch between oceanographic conditions that favor kelp survival and those that concentrate and distribute fish larvae. These results generally support the recruit-adult hypothesis: kelp bass populations are limited by recruitment at low recruit densities but by density-dependent competition for food resources and/or predator refuges at high recruit densities. At the same time, spatial variation in kelp abundance produced substantial spatiotemporal heterogeneity in kelp bass demographics, which argues for a multispecies, metacommunity approach to predicting kelp bass dynamics.     

Effects of Fishing on the Ecosystem Structure of Coral Reefs

Overfishing is considered one of the three most significant threats to coral reef ecosystems. Exponentially increasing human populations in the tropics have placed enormous demands upon reefs as a food source. At high intensities, termed ecosystem or Malthusian overfishing, fishing causes major direct and indirect effects on the community structure of fishes and other organisms. It reduces species diversity and leads to local extinctions not only of target species but also of other species not fished directly. Conceivably it could also lead to global extinctions. Loss of keystone species, such as predators of echinoderms, through fishing, can lead to major effects on reef processes, such as accretion of calcium carbonate. Ultimately, sustained heavy fishing may lead to loss of entire functional groups of species, resulting in impairment of the potentially important ecosystem functions provided by those groups. Overfishing has been shown to interact with other agents of disturbance to reduce the ability of reefs to recover from natural occurrences such as hurricanes. Effective management of fishing will require a deeper understanding of the effects of exploitation than we now possess. Research initiatives are underway to examine the responses of fish populations to fishing, generally responses to protection from fishing. There is, however, an urgent need to look beyond fish communities and to consider the entire reef ecosystem. Studies that integrate population and community biology with ecosystem processes will provide a much better understanding of the effects of biodiversity loss on reef function and will improve our ability to manage these complex systems.     

Effects of Aquarium Collectors on Coral Reef Fishes in Kona, Hawaii

Abstract:   No previous studies have conclusively documented the magnitude of the effect of aquarium collecting on natural populations. In Hawaii concern over the effects on reef fish populations of collecting for the aquarium trade began in the early 1970s, primarily in response to multiple-use conflicts between aquarium-fish collectors and recreational dive-tour operators. In 1997–1998 we used a paired control-impact design to estimate the effect of aquarium collectors. We compared differences in fish abundance along visual belt transects between collection sites, where collecting was known to occur, and control sites, where collecting was prohibited. To test the assumptions of our observational design, we surveyed a combination of species captured by aquarium collectors and those not captured. The extent of bleaching, broken coral, and coral cover was also surveyed. Seven of the 10 aquarium species surveyed were significantly reduced by collecting. The abundance of aquarium fish at collection sites ranged from 38% lower ( Chaetodon multicinctus ) to 75% lower ( C. quadrimaculatus ) than that at control sites. In contrast, only two of the nonaquarium species displayed a significant collection effect. There were no significant differences in damaged coral between control and collection sites to indicate the presence of destructive fishing practices. In addition, there were no increases in the abundance of macroalgae where the abundance of herbivores was reduced by aquarium collecting. Although our results suggest that aquarium collectors have a significant effect on the abundance of targeted aquarium fishes, better knowledge of the intensity and location of collecting activities is required to make a rigorous assessment of the effects of collecting on nearshore fish populations. Several lines of evidence suggest that the current system of catch reporting underestimates actual removals.     

Contrasting effects of habitat loss and fragmentation on coral-associated reef fishes

Disturbance can result in the fragmentation and/or loss of suitable habitat, both of which can have important consequences for survival, species interactions, and resulting patterns of local diversity. However, effects of habitat loss and fragmentation are typically confounded during disturbance events, and previous attempts to determine their relative significance have proved ineffective. Here we experimentally manipulated live coral habitats to examine the potential independent and interactive effects of habitat loss and fragmentation on survival, abundance, and species richness of recruitment-stage, coral-associated reef fishes. Loss of 75% of live coral from experimental reefs resulted in low survival of a coral-associated damselfish and low abundance and richness of other recruits 16 weeks after habitat manipulations. In contrast, fragmentation had positive effects on damselfish survival and resulted in greater abundance and species richness of other recruits. We hypothesize that spacing of habitat through fragmentation weakens competition within and among species. Comparison of effect sizes over the course of the study period revealed that, in the first six weeks following habitat manipulations, the positive effects of fragmentation were at least four times stronger than the effects of habitat loss. This initial positive effect of fragmentation attenuated considerably after 16 weeks, whereas the negative effects of habitat loss increased in strength over time. There was little indication that the amount of habitat influenced the magnitude of the habitat fragmentation effect. Numerous studies have reported dramatic declines in coral reef fish abundance and diversity in response to disturbances that cause the loss and fragmentation of coral habitats. Our results suggest that these declines occur as a result of habitat loss, not habitat fragmentation. Positive fragmentation effects may actually buffer against the negative effects of habitat loss and contribute to the resistance of reef fish populations to declines in coral cover.