Predation, habitat complexity, and variation in density-dependent mortality of temperate reef fishes

Density dependence in demographic rates can strongly affect the dynamics of populations. However, the mechanisms generating density dependence (e.g., predation) are also dynamic processes and may be influenced by local conditions. Understanding the manner in which local habitat features affect the occurrence and/or strength of density dependence will increase our understanding of population dynamics in heterogeneous environments. In this study I conducted two separate field experiments to investigate how local predator density and habitat complexity affect the occurrence and form of density-dependent mortality of juvenile rockfishes (Sebastes spp.). I also used yearly censuses of rockfish populations on nearshore reefs throughout central California to evaluate mortality of juvenile rockfish at large spatial scales. Manipulations of predators (juvenile bocaccio, S. paucispinus) and prey (kelp, gopher, and black-and-yellow [KGB] rockfish, Sebastes spp.) demonstrated that increasing the density of predators altered their functional response and thus altered patterns of density dependence in mortality of their prey. At low densities of predators, the number of prey consumed per predator was a decelerating function, and mortality of prey was inversely density dependent. However, at high densities of predators, the number of prey killed per predator became an accelerating response, and prey mortality was directly density dependent. Results of field experiments and large-scale surveys both indicated that the strength of density-dependent mortality may also be affected by the structural complexity of the habitat. In small-scale field experiments, increased habitat complexity increased the strength of density-dependent mortality. However, at large scales, increasing complexity resulted in a decrease in the strength of density dependence. I suggest that these differences resulted from scale-dependent changes in the predatory response that generated mortality. Whether increased habitat complexity leads to an increase or a decrease in the strength of density-dependent mortality may depend on how specific predatory responses (e.g., functional or aggregative) are altered by habitat complexity. Overall, the findings of this study suggest that rates of demographic density dependence and the resulting dynamics of local populations may largely depend upon attributes of the local habitat.     

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.     

Associations between the abundance of piscivorous fishes and their prey on coral reefs: implications for prey-fish mortality

Few studies have examined predator-prey relationships in diverse communities such as those found on coral reefs. Here we examined patterns of spatial and temporal association between the local abundance of predator and prey fishes at Lizard Island on the Great Barrier Reef, Australia. We predicted that the nature of this association would have implications for patterns of prey-fish mortality. Strong positive relationships between prey and piscivore abundance were found throughout the study. Greater densities of predators and of prey were found on patch-reef habitats, compared with contiguous reef-slope habitats. Declines in prey-fish abundance on patch reefs were density-dependent and correlated with the densities of predators. The relative roles of recruitment and piscivore movement in determining patterns of predator and prey abundance were assessed from surveys of recruit densities and an intensive programme of tagging two species of rock-cod, Cephalopholis cyanostigma and C. boenak (Serranidae), over 2 years. Patterns of recruitment explained little of the variation in the abundance and distribution of piscivorous fish. If movement explains large-scale patterns of distribution, this was not evident from the tagging study. The two rock-cod species were highly sedentary, with individuals on patch reefs seldom moving among reefs. Individuals on reef slopes were also highly site-attached, although they moved greater distances than those on patch reefs. Although the mechanisms responsible remain to be determined, this study demonstrated strong associations between the abundance of piscivorous fish and their prey on coral reefs. This relationship appeared to be an important factor in producing density-dependent declines in the abundance of prey. Received: 30 April 2000 / Accepted: 22 September 2000     

Safety in numbers and the spatial scaling of density-dependent mortality in a coral reef fish

In coral reef fishes, density-dependent population regulation is commonly mediated via predation on juveniles that have recently settled from the plankton. All else being equal, strong density-dependent mortality should select against the formation of high-density aggregations, yet the juveniles of many reef fishes aggregate. In light of this apparent contradiction, we hypothesized that the form and intensity of density dependence vary with the spatial scale of measurement. Individual groups might enjoy safety in numbers, but predators could still produce density-dependent mortality at larger spatial scales. We investigated this possibility using recently settled juvenile bluehead wrasse, Thalassoma bifasciatum, a small, aggregating reef fish. An initial caging experiment demonstrated that juvenile bluehead wrasse settlers suffer high predation, and spatial settlement patterns indicated that bluehead wrasse juveniles preferentially settle in groups, although they are also found singly. We then monitored the mortality of recently settled juveniles at two spatial scales: microsites, occupied by individual fish or groups of fish and separated by centimeters, and sites, consisting of approximately 2400-m2 areas of reef and separated by kilometers. At the microsite scale, we measured group size and effective population density independently and found that per capita mortality decreased with group size but was not related to density. At the larger spatial scale, however, per capita mortality increased with settler density. This shift in the form of density dependence with spatial scale could reconcile the existence of small-scale aggregative behavior typical of many reef fishes with the population-scale density dependence that is essential to population stability and persistence.     

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.     

Effects of predation and habitat structure on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Caridea) on temperate rocky reefs

Human disturbances, such as overfishing, may disrupt predator–prey interactions and modify food webs. Underwater surveys were carried out at six shallow-water reef barrens in temperate waters of northern-central Chile from October to December 2010 to describe the effects of predation, habitat complexity (low, medium and high) and refuge availability on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Rhynchocinetidae), an important mesoconsumer on subtidal hard substrata. Three sites were within managed (restricted access) areas for fishermen, and three were unmanaged (open-access). Field observations and tethering experiments were conducted to examine the relationship between fish and shrimp abundances, and the relative predation rates on shrimps. Direct effects of predation on R. typus body-size distribution were examined from shrimps collected in the field and fish stomachs. The presence and the abundance of R. typus increased with habitat reef complexity and refuge availability. Shrimp abundance was negatively related to fish abundance in managed areas, but not in open-access areas, where shrimp densities were the highest. Also, predation rates and body-size distribution of shrimps were unrelated, although fish consumed more large shrimps than should be expected from their distribution in the field. R. typus occurred most often in shelters with wide openings, offering limited protection against predators, but providing potential aggregation sites for shrimps. Overall, direct effects of predation on shrimp densities and population structure were weak, but indirect effects on shrimp distribution within reefs appear to have been mediated through behavioural responses. Our study highlights the need to assess both numerical and behavioural responses of prey to determine the effects of predator loss on mesoconsumer populations.     

Consumptive and nonconsumptive effects of predators on metacommunities of competing prey

Although predators affect prey both via consumption and by changing prey migration behavior, the interplay between these two effects is rarely incorporated into spatial models of predator-prey dynamics and competition among prey. We develop a model where generalist predators have consumptive effects (i.e., altering the likelihood of local prey extinction) as well as nonconsumptive effects (altering the likelihood of colonization) on spatially separated prey populations (metapopulations). We then extend this model to explore the effects of predators on competition among prey. We find that generalist predators can promote persistence of prey metapopulations by promoting prey colonization, but predators can also hasten system-wide extinction by either increasing local extinction or reducing prey migration. By altering rates of prey migration, predators in one location can exert remote control over prey dynamics in another location via predator-mediated changes in prey flux. Thus, the effect of predators may extend well beyond the proportion of patches they visit. In the context of prey metacommunities, predator-mediated shifts in prey migration and mortality can shift the competition-colonization trade-off among competing prey, leading to changes in the prey community as well as changes in the susceptibility of prey species to habitat loss. Consequently, native prey communities may be susceptible to invasion not only by exotic prey species that experience reduced amounts of mortality from resident predators, but also by exotic prey species that exhibit strong dispersal in response to generalist native predators. Ultimately, our work suggests that the consumptive and nonconsumptive effects of generalist predators may have strong, yet potentially cryptic, effects on competing prey capable of mediating coexistence, fostering invasion, and interacting with anthropogenic habitat alteration.     

Food webs and intraguild predation: community interactions of a native mesocarnivore

Trophic level interactions between predators create complex relationships such as intraguild predation. Theoretical research has predicted two possible paths to stability in intraguild systems: intermediate predators either outcompete higher-order predators for shared resources or select habitat based on security. The effects of intraguild predation on intermediate mammalian predators such as swift foxes (Vulpes velox) are not well understood. We examined the relationships between swift foxes and both their predators and prey, as well the effect of vegetation structure on swift fox-coyote (Canis latrans) interactions, between August 2001 and August 2004. In a natural experiment created by the Pinon Canyon Maneuver Site in southeastern Colorado, USA, we documented swift fox survival and density in a variety of landscapes and compared these parameters in relation to prey availability, coyote abundance, and vegetation structure. Swift fox density varied significantly between study sites, while survival did not. Coyote abundance was positively related to the basal prey species and vegetation structure, while swift fox density was negatively related to coyote abundance, basal prey species, and vegetation structure. Our results support the prediction that, under intraguild predation in terrestrial systems, top predator distribution matches resource availability (resource match), while intermediate predator distribution inversely matches predation risk (safety match). While predation by coyotes may be the specific cause of swift fox mortality in this system, the more general mechanism appears to be exposure to predation moderated by shrub density.     

Propagule redirection: habitat availability reduces colonization and increases recruitment in reef fishes

Increased habitat availability or quality can alter production of habitat-dependent organisms in two contrasting ways: (1) by enhancing input of new colonists to the new sites (the Field-of-Dreams Hypothesis); and (2) by drawing colonists away from existing sites (the Propagule Redirection Hypothesis), and thus reducing the deleterious effects of density. We conducted a field experiment on coral reef fishes in Moorea, French Polynesia, to quantify how differing levels of habitat availability (controlling for quality) increased and/or redirected colonizing larval fish. Focal reefs without neighboring reefs received two to four times more settlers than reefs with adjacent habitat, demonstrating that increased habitat redirected larval fish. At the scale of the entire reef array, total colonization increased 1.3-fold in response to a sixfold increase in reef area (and a 2.75-fold increase in adjusted habitat availability). Thus, propagules were both increased and redirected, a result midway between the Field-of-Dreams and Propagule Redirection Hypotheses. A recruitment model using our data and field estimates of density-dependent recruitment predicts that habitat addition increases recruitment primarily by ameliorating the negative effects of competition at existing sites rather than increasing colonization at the new sites per se. Understanding long-term implications of these effects depends upon the interplay among habitat dynamics, population connectivity, colonization dynamics, and density dependence.     

Predicting local population distributions around a central shelter based on a predation risk-growth trade-off

Animals face trade-offs between predation risk and foraging success depending on their location in the landscape; for example, individuals that remain near a common shelter may be safe from predation but incur stronger competition for resources. Despite a long tradition of theoretical exploration of the relationships among foraging success, conspecific competition, predation risk, and population distribution in a heterogeneous environment, the scenario we describe here has not been explored theoretically. We construct a model of habitat use rules to predict the distribution of a local population (prey sharing a common shelter and foraging across surrounding habitats). Our model describes realized habitat quality as a ratio of density- and location-dependent mortality to density-dependent growth. We explore how the prey distribution around a shelter is expected to change as the parameters governing the strength of density dependence, landscape characteristics, and local abundance vary. Within the range of parameters where prey spend some time away from shelter but remain site-attached, the prey density decreases away from shelter. As the distance at which prey react to predators increases, the population range generally increases. At intermediate reaction distances, however, increases in the reaction distance lead to decreases in the maximum foraging distance because of increased evenness in the population distribution. As total abundance increases, the population range increases, average population density increases, and realized quality decreases. The magnitude of these changes differs in, for example, ‘high-’ and ‘low-visibility’ landscapes where prey can detect predators at different distances.     

Density dependence in marine fish populations revealed at small and large spatial scales

Experimental manipulation of population density has frequently been used to demonstrate demographic density dependence. However, such studies are usually small scale and typically provide evidence of spatial (within-generation) density dependence. It is often unclear whether small-scale, experimental tests of spatial density dependence will accurately describe temporal (between-generation) density dependence required for population regulation. Understanding the mechanisms generating density dependence may provide a link between spatial experiments and temporal regulation of populations. In this study, I manipulated the density of recently settled kelp rockfish (Sebastes atrovirens) in both the presence and absence of predators to test for density-dependent mortality and whether predation was the mechanism responsible. I also examined mortality of rockfish cohorts within kelp beds throughout central California to evaluate temporal (between-generation) density dependence in mortality. Experiments suggested that short-term behavioral responses of predators and/or a shortage of prey refuges caused spatial density dependence. Temporal density dependence in mortality was also detected at larger spatial scales for several species of rockfish. It is likely that short-term responses of predators generated both spatial and temporal density dependence in mortality. Spatial experiments that describe the causal mechanisms generating density dependence may therefore be valuable in describing temporal density dependence and population regulation.     

Patch use and vigilance by sympatric lemmings in predator and competitor-driven landscapes of fear

Prey living in risky environments can adopt a variety of behavioral tactics to reduce predation risk. In systems where predators regulate prey abundance, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, patterns of habitat use also reflect interspecific competition over habitat. Collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings represent such a system and possess distinct upland tundra versus mesic meadow habitat preferences consistent with interspecific competition. Yet, we do not know whether this habitat preference might also reflect differences in predation risk or whether the two species differ in their behavioral tactics used to avoid predation. We performed experiments where we manipulated putative predation risk perceived by lemmings by increasing protective cover in upland and meadow habitats while we recorded lemming activity and behavior. Both lemming species preferentially used cover more than open patches, but Dicrostonyx was more vigilant than Lemmus. Both species also constrained their activity to protective patches in upland and meadow habitats, but during different periods of the day. Use of cover and vigilance were independent of habitat, suggesting that both species live in a fearsome but flattened landscape of fear at Walker Bay (Nunavut, Canada), and that their habitat preference is a consequence of competition rather than predation risk. Future studies aiming to map the contours of fear in multi-prey–predator systems should consider how predation and competition interact to modify prey species’ habitat preference, patch use, and vigilance.     

Female nest defense in a coral-reef fish, Dascyllus albisella, with uniparental male care

Intraspecific variation in the patterns of parental care has been observed in a variety of animals; however, the possibility of parental care by a non-caregiving parent of uniparental species has not been thoroughly explored. In the coral-reef damselfish, Dascyllus albisella, only males normally exhibit parental care. In this study, we examined the response of females of this species to egg predators after experimental male removal and an elevated level of egg predation, at two small patch reefs (reefs 1 and 2) in Hawaii. We tested theoretical expectations that a nest was defended only by females which had spawned in the nest, and that larger females had a higher likelihood of defense than smaller females. A nest was defended against egg predators more frequently by females that had spawned in that nest than would be expected by chance. Not all females that had spawned in a given nest participated in defense. There was a positive association between female body length and the likelihood of defense at reef 2, but not at reef 1. Within a set of females that had spawned in the same nest during the same nesting cycle, defending females had larger body lengths than non-defending females at reef 2 but not at reef 1. Lack of association between female size and likelihood of defense at reef 1 was unexpected, but may correlate with the smaller average female size and smaller size differences among females on that reef.     

Effects of sheltering fish on growth of their host corals

Stony corals are the foundation species of tropical reefs, and their structures can harbor a diverse range of mutualist taxa that can confer important benefits, including provision of nutrients. Prominent among the associates of branching coral in the genus Pocillopora are groups of zooplanktivorous damselfishes that take refuge in the coral to avoid their predators. In field and laboratory experiments, we explored the effects of colonies of resident damselfishes on growth of their host corals. Laboratory studies revealed a positive relationship between biomass of fish and output of ammonium. In the field, levels of ammonium were significantly elevated in the water surrounding the branches of Pocillopora occupied by colonies of damselfish, particularly in time periods following active feeding by the fish. Experimental manipulation of the presence of fish on host corals during a month-long field experiment revealed that corals hosting fish grew significantly more than those that lacked fish, and coral growth was positively correlated with the biomass of resident fish. The Pocillopora colonies in the field experiment varied in the degree of openness of their branching structure, and dye studies indicated that this affected their ability to retain waterborne nutrients. Together with biomass of resident fish, colony openness explained 76% of the variation in coral growth rate during the experiment. Corals can exhibit considerable morphological variability, and mutualistic fish respond to colony architecture during habitat selection, with some species preferring more open-branched forms. This makes it likely that corals may face tradeoffs in attracting resident fish and in retaining the nutrients they provide.     

Estuarine Vegetated Habitats as Corridors for Predator Movements

Abstract: The spatial proximity of one habitat to another can strongly influence population and community dynamics. We investigated whether the proximity of intertidal oyster reefs to vegetated estuarine habitats, salt marshes, and seagrass beds, affects the abundance and community structure of benthic macroinvertebrates on reefs and predator-prey interactions between mobile predators and bivalves living on reefs. Benthic macroinvertebrate abundance was highest on reefs spatially separated from salt marshes. Macroinvertebrate species richness was highest on reefs separated from both salt marshes and seagrass beds. Comparisons of predation on juvenile bivalves transplanted to reefs for 7–12 days indicated that survivorship of clams was greatest on reefs spatially separated from both salt marshes and seagrass beds, whereas reef proximity to vegetated habitats did not affect the survivorship of oysters. The foraging behavior of blue crabs may explain patterns of macroinvertebrate abundance and clam survivorship among reefs with different proximity to vegetated habitats. In experiments conducted in 30-m² field enclosures, blue crabs had higher predation rates on hard clams transplanted onto artificial reefs adjacent to salt marshes or seagrass beds than on reefs separated from both habitats by unvegetated sand bottom. Thus, vegetated habitats appeared to act as corridors by facilitating the access of blue crabs to oyster reefs and enhancing the intensity of blue crab predation. Such an understanding of the effects of landscape characteristics of estuarine habitats on their value as habitats for estuarine organisms can be used to predict the consequences of habitat fragmentation on ecosystem function and to improve strategies for habitat and species conservation and restoration.     

Use of pelagic prey subsidies by demersal predators in rocky reefs: insight from movement patterns of lingcod

Allochthonous subsidies of energy and nutrients can affect community structure in patchy marine habitats, including rocky reefs, and their ecological consequences may depend on the mechanism of energy transfer. Lingcod (Ophiodon elongatus) are demersal predators that trophically link nearshore rocky reefs with offshore pelagic habitats through consumption of pelagic fishes. We quantified lingcod habitat use and movement patterns to make inferences about the temporal and spatial conditions under which lingcod may acquire pelagic prey. Lingcod maintained small home ranges (21,272 ± 13,630 m²) within a rocky reef in the San Juan Archipelago, Washington; eight of nine individuals used rocky habitat exclusively. Depths occupied by lingcod (0–50 m) coincided with pelagic fish distribution on the rocky reef; however, diel patterns in lingcod activity varied inversely with occurrence of pelagic fishes on the reef. Our findings suggest that the pelagic subsidy to lingcod is not strongly mediated through directed off-reef foraging by lingcod.     

Prevalence and distribution patterns of tumors in bicolor damselfish (Pomacentrus partitus) on South Florida reefs

Damselfish neurofibromatosis (DNF) is a cancer affecting bicolor damselfish (Pomacentrus partitus) on reefs in South Florida, USA. Bicolor damselfish exhibiting DNF were observed on 18 of 19 reefs surveyed, with prevalence rates varying from 0.4 to 23.8% of the adult fish in these populations. These rates were very stable over 5 to 9 yr periods (1981 to 1989) on the two reefs monitored. High disease prevalence was associated with a high population density of large size-class fish and a population structure dominated by large fish. These patterns were observed both between and within reef populations. The distribution of cases within reefs was clustered rather than uniform or random. These findings are consistent with the hypothesis that DNF is distributed primarily via fish to fish (horizontal) transmission of an infectious agent. Disease patterns are not consistent with what would be expected to result from inherited (or vertically transmitted) or water-borne agents.     

Ecology of Conus on eastern Indian Ocean fringing reefs: Diversity of species and resource utilization

The most diverse assemblages of the genus Conus known occur on fringing coral reefs in Thailand and Indonesia. As many as 27 congeneric species of these gastropods inhabit a single reef; in all, we examined 1,350 individuals of 48 species. Several attributes of the populations we observed conform to expectations of a model of ecological characteristics of bench and reef Conus proposed by Kohn (1971a). Number of species (S) averaged 15, and species diversity (H″) averaged 2.3 in the most heterogeneous habitat type — topographically complex, subtidal reef platforms (Type III habitat). Both species richness and evenness of distribution of individuals among species contribute strongly to H″. Fewer congeners and greater numerical dominance by single species characterize more homogeneous habitats. On subtidal reef platforms with large areas of sand substrate and less coral limestone (Type I–III habitat), mean values were S=10, and H″=1.6. In the one intensively studied, truncated reef-limestone platform (Type II–III intermediate habitat), S=13 and H″=1.4. Summed population density of all Conus species in Type III and I–III habitats is similar (0.02 to 0.05 individuals /m²) and comparable to estimates from similar habitats elsewhere in the Indo-West Pacific region. Mean density (0.7/m²) and other population attributes in Type II–III habitat more closely resemble those of Type II than Type III habitats in general. We combined analysis of species diversity and other attributes of assemblages in habitats of different environmental complexity with analysis of microhabitat and food-resource utilization, in order to demonstrate the extent to which specialization on different resources occurs in assemblages differing in diversity and habitat type. In the habitats studied, co-occurring species of Conus specialized to a greater extent on different prey species than on different microhabitat patches, but degree of microhabitat specialization was greater than in similarly complex habitats with assemblages of lower diversity elsewhere in the Indo-West Pacific region. While most Conus species preyed primarily on a different species or higher taxon of polychaetes, diets are not more specialized or dissimilar than in similar habitats elsewhere. Degree of specialization on different prey is not correlated with Conus species diversity in the different types of habitats studied. The data lead to the conclusion that differential predation is as important — and differential microhabitat utilization is more important — in permitting coexistence of potentially competing congeners, compared with conditions in habitats of comparable heterogeneity that support fewer congeners farther from the center of the Indo-West Pacific region. Pairwise comparisons of congeners indicate that many species pairs have low or no overlap in both microhabitat and food utilization. Members of species pairs with high overlap in microhabitat utilization typically eat different prey organisms, and those with similar diets typically occupy different habitats or microhabitats. This applies to molluscivorous as well as vermivorous species. Information on the diets of 11 species is reported here for the first time. Of 48 Indo-West Pacific Conus species whose food is now known, 35 prey on polychaetes, 2 on enteropneusts, 6 on gastropods, and 5 on fishes. Vermivorous Conus prey on relatively few of the polychaete species present in the environments. Species eaten represent only 12% of a total estimated polychaete population density of 27,000 individuals /m². Certain very abundant polychaetes may be protected from predation by Conus by their small size, others by their long tubes. Two new aspects of size-selective predation by Conus are reported: (1) Although comparisons of predation rate with prey standing-crop suggest that food is plentiful, selective predation on the largest prey individuals present suggests that only small proportions of prey-species populations may have large enough body size to repay foraging effort by the Conus present; (2) composition of the diet changes qualitatively with increase in body size in several vermivorous Conus species; shifting by larger individuals to larger prey species could be documented in C. ebraeus.     

Antipredator responses to invasive lionfish, Pterois volitans: interspecific differences in cue utilization by two coral reef gobies

Understanding prey response to predators and their utilization of sensory cues to assess local predation risk is crucial in determining how predator avoidance strategies affect population demographics. This study examined the antipredator behaviors of two ecologically similar species of Caribbean coral reef fish, Coryphopterus glaucofraenum and Gnatholepis thompsoni, and characterized their responses to different reef predators. In laboratory assays, the two species of gobies were exposed to predator visual cues (native Nassau grouper predator vs. invasive lionfish predator), damage-released chemical cues from gobies, and combinations of these, along with appropriate controls. Behavioral responses indicate that the two prey species differ in their utilization of visual and chemical cues. Visual cues from predators were decisive for both species’ responses, demonstrating their relative importance in the sensory hierarchy, whereas damage-released cues were a source of information only for C. glaucofraenum. Both prey species could distinguish between native and invasive predators and subsequently altered their antipredator responses.     

Performance level and efficiency of two differing predator-avoidance strategies depend on nutritional state of the prey fish

Animal prey has developed a variety of behavioural strategies to avoid predation. Many fish species form shoals in the open water or seek refuge in structurally complex habitats. Since anti-predator strategies bear costs and are energy-demanding, we hypothesised that the nutritional state of prey should modify the performance level and efficiency of such strategies. In aquaria either containing or lacking a structured refuge habitat, well-fed or food-deprived juvenile roach (Rutilus rutilus) were exposed to an open-water predator (pikeperch, Sander lucioperca). Controls were run without predators. In the presence of the predator, roach enhanced the performance of the anti-predator strategy and increased the use of the refuge habitat whereby food-deprived roach were encountered more often in the structure than well-fed roach. Nonetheless more starved than well-fed roach were fed upon by the predator. In the treatments offering only open-water areas, roach always formed dense shoals in the presence of the predator. The shoal density, however, was lower in starved roach. Starving fish in shoals experienced the highest predation mortality across all experimental treatments. The experiment confirmed the plasticity of the anti-predator behaviour in roach and demonstrated that food deprivation diminished the efficiency of shoaling more strongly than the efficiency of hiding. The findings may be relevant to spatial distribution of prey and predator–prey interactions under natural conditions because when prey are confronted with phases of reduced resource availability, flexible anti-predator strategies may lead to dynamic habitat use patterns.