Consequences of patch reef spacing for density-dependent mortality of coral-reef fishes

The spatial configuration of habitat patches can profoundly affect a number of ecological interactions, including those between predators and prey. I examined the effects of reef spacing on predator-prey interactions within coral-reef fish assemblages in the Bahamas. Using manipulative field experiments, I determined that reef spacing influences whether and how density-dependent predation occurs. Mortality rates of juveniles of two ecologically dissimilar species (beaugregory damselfish and yellowhead wrasse) were similarly affected by reef spacing; for both species, mortality was density dependent on reef patches that were spatially isolated (separated by 50 m), and density independent on reef patches that were aggregated (separated by 5 m). A subsequent experiment with the damselfish demonstrated that a common resident predator (coney) caused a substantial proportion of the observed mortality, independent of reef spacing. Compared to isolated reefs, aggregated reefs were much more likely to be visited by transient predators (mostly yellowtail snappers), regardless of prey density, and on these reefs, mortality rates approached 100% for both prey species. Transient predators exhibited neither an aggregative response nor a type 3 functional response, and consequently were not the source of density dependence observed on the isolated reefs. These patterns suggest that resident predators caused density-dependent mortality in their prey through type 3 functional responses on all reefs, but on aggregated reefs, this density dependence was overwhelmed by high, density-independent mortality caused by transient predators. Thus, the spatial configuration of reef habitat affected both the magnitude of total predation and the existence of density-dependent mortality. The combined effects of the increasing fragmentation of coral reef habitats at numerous scales and global declines in predatory fish may have important consequences for the regulation of resident fish populations.     

Predation,recruitment and the dynamics of communities of coral-reef fishes

I investigated the ability of predators to influence the patterns of species richness and abundance of non-piscivorous fishes on small, artificial reefs replenished by natural recruitment. Periodic removal of predators effectively reduced the species richness and abundance of predators on removal reefs. The difference between the number of predators on control and removal reefs was greatest immediately following the removal of predators and attenuated between removals. During periods of recruitment, species richness and total abundance of recently-recruited, non-piscivorous fishes were generally greater on predator-removal reefs than on control reefs. Species richness and total abundance of resident non-piscivorous fishes were not affected by the removal of predators in the first year of the experiment. Both abundance and species richness of residents, however, were greater on the removal reefs during the second year of the experiment. The difference in the responses of the two age classes to the removal of predators suggests that predators may affect community patterns of older age classes through time-lagged effects on the survivorship of younger age classes. At the end of the experiment, species richness was positively related to abundance for recruits and residents. The effects of removing piscivorous fishes on the abundance of non-piscivorous fishes were similar for species considered separately. A greater number of species of recruit and resident fishes were more abundant on reefs from which predators had been removed. These data suggest that predators can play an important role in structuring communities of fishes on coral reefs.     

Variability in abundances of fishes associated with seagrass habitats in relation to diets of predatory fishes

The spatial, diel and tidal variability in the abundance of piscivorous fishes and their teleost prey, and the dietary composition of predatory fishes were investigated in beds of Heterozostera tasmanica within Port Phillip Bay, Australia, from September 1997 to February 1998. Predatory and prey fish assemblages were sampled from beds of H. tasmanica at three locations during each combination of diel (day and night) and tidal (high and low) cycles. Pelagic and benthic crustaceans represented >60% by abundance of the diets of all predatory fishes. Seven species, 54% of all predatory fishes, were piscivorous. These piscivores consumed individuals from seven families, 36.8% of the fish families being associated with seagrass. Western Australian salmon, Arripis truttacea (Arripidae) (n = 174) and yank flathead, Platycephalus speculator (Platycephalidae) (n = 46) were the most abundant piscivores. A. truttacea consumed larval/post-larval atherinids, gobiids and sillaginids. P. speculator consumed late-juvenile/adult atherinids, clinids and gobiids. While the abundances of piscivores varied between locations (P < 0.001) and diel periods (P = 0.028), the relative differences in piscivore abundance between sites and diel periods were not consistent between tides. The abundances of A. truttacea varied in a complex way amongst sites, diel period and tidal cycle, as shown by a three-way interaction between these factors (P = 0.026). Only during diurnal periods at St. Leonards was the abundance of A. truttacea significantly higher during high than low tides (P < 0.001). During the other diel periods at each site, the abundance of A. truttacea did not vary. P. speculator was significantly more abundant nocturnally (P = 0.017). The abundance of small (prey) fishes varied significantly amongst sites (P < 0.001). During the day, the abundance of small fishes did not vary between high and low tides (P = 0.185), but their nocturnal abundance was greater during low tide (P < 0.001). Atherinids (n = 1732) and sillaginids (n = 1623) were the most abundant families of small fishes. Atherinids were significantly more abundant nocturnally (P = 0.005) and during low tides (P = 0.029), and varied significantly amongst sites (P < 0.001). Sillaginids varied significantly only amongst sites (P < 0.001). Seagrass beds provide a foraging habitat for a diverse assemblage of predatory fishes, many of which are piscivorous. Anti-predator behaviour and amongst-location variability in abundances of piscivorous fishes may explain some of the diel and tidal, and broad-scale spatial patterns in small-fish abundances. Received: 23 July 1999 / Accepted: 18 January 2000     

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.     

Habitat patchiness and predation modify the distribution of a coral-dwelling damselfish

Fish abundance is often better predicted by microhabitat variables on continuous reefs than on isolated patch reefs. Although this was suggested to stem from reduced post-recruitment relocation, this has not been shown experimentally. We found the relationship between the presence of a coral-dwelling fish, Dascyllus marginatus, and the size of its coral host to differ between corals on continuous reefs and the sparsely distributed corals on sandy bottoms. Empty transplanted corals were colonized exclusively by new recruits when on the sandy bottom, and both by new recruits and post-recruitment dispersal of adults when on the continuous reef. New recruits settled predominantly into small corals, although analyses of recruitment patterns were confounded by low recruitment in the studied years. Both tank experiments and field survey data suggest that the presence of recruits in small corals is at least partially driven by predation by the dottyback, Pseudochromis olivaceus, which lives predominantly in large corals within both habitats. Consequently, we suggest that the relationship between fish presence and coral size differs between the habitats due to coral size dependent predation on recruits and variability in the importance of direct recruitment to replenish fish populations.     

Discarded queen conch (<Emphasis Type="Italic">Strombus gigas</Emphasis>) shells as shelter sites for fish

Within the Caribbean millions of queen conch (Strombus gigas Linnaeus) are harvested each year and shells discarded randomly or as middens. Fish use of discarded conch shells was investigated in four different habitat types: sand, seagrass beds, mangrove forests, and coral reefs. The study was carried out in the waters off South Caicos, Turks and Caicos Islands (TCI), between October 2003 and January 2004. The density of discarded shells was greatest near coral reefs; however, the percentage of shells occupied by adult fish was higher in isolated shells on sand and in mangrove habitats. Juvenile fish also showed a preference for sheltering in conch shells relative to other microhabitat types on sandy plains and in mangrove and seagrass habitats. Differences in use of single shells by fish in different habitats were attributed to differences in piscivore abundance and habitat complexity. Although not all isolated shells were occupied by fish, all conch middens deposited by fishermen had fish inhabitants. Examination of fish use of conch middens in three habitat types and conch piles of one, three, and five shells constructed on sand found both fish diversity and abundance increased on conch middens of increasing size. This study suggests that disposal of conch shells as large middens in habitats of low complexity will increase the amount of shelter present and may enhance fish populations in these habitats.Communicated by J.P. Grassle, New Brunswick     

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.     

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.     

The paradox of risk assessment: comparing responses of fathead minnows to capture-released and diet-released alarm cues from two different predators

Summary. Many aquatic prey are known to use chemical alarm cues to assess their risk of predation. In fishes, such alarm cues can be released either through damage of the epidermis during a predatory attack (capture-released) or through release from the predator feces (diet-released). In our study, we compared the importance of capture- versus diet-released alarm cues in risk assessment by fathead minnows (Pimephales promelas) that were na?ve to fish predators. We utilized two different fish predators: a specialized piscivore, the northern pike (Esox lucius) and a generalist predator, the brook trout (Salvelinus fontinalis). Handling time of pike consuming minnows was much shorter than for trout consuming minnows, likely resulting in less epidermal damage to the minnows during attacks by pike. In accordance with this, minnows showed a less intense antipredator response to capture-released cues from pike than capture-released cues from trout. This represents a paradox in risk assessment for the minnows as they respond to the specialized piscivore, the more dangerous predator, with a less intense antipredator response. In contrast, the minnows showed a stronger antipredator response to the specialized piscivore than to the generalist when given diet cues. This work highlights the need for researchers to carefully consider the nature of the information available to prey in risk assessment.     

In situ swimming and settlement behaviour of larvae of an Indo-Pacific coral-reef fish, the coral trout Plectropomus leopardus (Pisces: Serranidae)

Late larvae of the serranid coral trout Plectropomus leopardus (Lacepède), captured in light traps, were released during the day both in open water and adjacent to two reefs, and their behaviour was observed by divers at Lizard Island, northern Great Barrier Reef. Coral trout larvae (n = 110) were present in light-trap catches from 18 November to 3 December 1997, including new moon (30 November). The swimming speed of larvae in open water or when swimming away from reefs was significantly greater (mean 17.9 cm s⁻¹) than the speed of larvae swimming towards or over reefs (mean 7.2 cm s⁻¹). Near reefs, larvae swam at average depths of 2.7 to 4.2 m, avoiding 0 to 2 m. In open water, swimming depth varied with location: larvae >1 km east of Lizard Island swam steeply downward to >20 m in 2 to 4 min; larvae >1 km west oscillated between 2.6 and 13 m; larvae 100 to 200 m east of Lizard Island oscillated between 0.8 and 15 m. Nearly all larvae swam directionally in open water and near reefs. In open water, the average swimming direction of all larvae was towards the island, and 80% (4 of 5) swam directionally (p < 0.05, Rayleigh's test). Larvae swam directionally over the reef while looking for settlement sites. The frequency of behaviours by larvae differed between two reefs of different exposure and morphology. Depending on site, 26 to 32% of larvae released adjacent to reefs swam to open water: of these, some initially swam towards or over the reef before swimming offshore. In some cases, offshore-swimming seemed to be due to the presence of predators, but usually no obvious cause was observed. Depending on the reef, 49 to 64% of the larvae settled. Non-predatory reef residents aggressively approached 19% of settlers. Between 5 and 17% of the larvae were eaten while approaching the reef or attempting to settle, primarily by lizardfishes but also by wrasses, groupers and snappers. A higher percentage of larvae settled in the second week of our study than in the first. Average time to settlement was short (138 s ± 33 SE), but some larvae took up to 15 min to settle. Average settlement depth was 7.5 to 9.9 m, and differed between locations. No settlement took place on reef flats or at depths <4.2 m. Larvae did not appear to be selective about settlement substrate, but settled most frequently on live and dead hard coral. Late-stage larvae of coral trout are capable swimmers with considerable control over speed, depth and direction. Habitat selection, avoidance of predators and settlement seem to rely on vision. Received: 7 July 1998 / Accepted: 26 January 1999     

Impact of conservation areas on trophic interactions between apex predators and herbivores on coral reefs

Apex predators are declining at alarming rates due to exploitation by humans, but we have yet to fully discern the impacts of apex predator loss on ecosystem function. In a management context, it is critically important to clarify the role apex predators play in structuring populations of lower trophic levels. Thus, we examined the top‐down influence of reef sharks (an apex predator on coral reefs) and mesopredators on large‐bodied herbivores. We measured the abundance, size structure, and biomass of apex predators, mesopredators, and herbivores across fished, no‐take, and no‐entry management zones in the Great Barrier Reef Marine Park, Australia. Shark abundance and mesopredator size and biomass were higher in no‐entry zones than in fished and no‐take zones, which indicates the viability of strictly enforced human exclusion areas as tools for the conservation of predator communities. Changes in predator populations due to protection in no‐entry zones did not have a discernible influence on the density, size, or biomass of different functional groups of herbivorous fishes. The lack of a relationship between predators and herbivores suggests that top‐down forces may not play a strong role in regulating large‐bodied herbivorous coral reef fish populations. Given this inconsistency with traditional ecological theories of trophic cascades, trophic structures on coral reefs may need to be reassessed to enable the establishment of appropriate and effective management regimes. El Impacto de las Áreas de Conservación sobre las Interacciones Tróficas entre los Depredadores Dominantes y los Herbívoros en los Arrecifes de Coral     

Effects of herbivory on zonation of Sargassum spp. within fringing reefs of the central Great Barrier Reef

A combination of small-scale transplants and herbivore exclusion was used to test the importance of herbivory, physiological tolerance limits, and recruitment and dispersal in regulating the distribution and abundance of the genus Sargassum on two nearshore fringing reefs of the central Great Barrier Reef, during 1992/1993. Sargassum (predominantly S. oligocystum and S. tenerrimum) were transplanted from reef-flat zones where they normally grow, to a seaward coral zone where they are not normally found. At Great Palm Island, coral-zone transplants only survived if protected from herbivores. At Brook Island, survival of uncaged coral-zone transplants was more variable but not significantly lower than plants returned to the Sargassum zone. Thus herbivory may be a major cause of the zonation patterns of adult Sargassum on these fringing reefs, but the importance of this factor varies between and within reefs. Since protected Sargassum survived and grew for up to 6 mo in the coral zone, the adult algae are not physiologically limited by any physical or chemical differences between zones. However, Sargassum recruitment to the coral zone was very low (mean 2.7 recruits m⁻² over 13 mo), and was not significantly affected by herbivores. Since rates of herbivory were relatively slow, effective exclusion of Sargassum from the coral zone by herbivores may depend on low recruitment of the algae. In a broader context, the distribution of Sargassum may depend on the combined spatial patterns of herbivory and recruitment. Received: 24 January 1997 / Accepted: 12 May 1997     

Effects of combining an intraguild predator with a cannibalistic intermediate predator on a species-level trophic cascade

A greater diversity of natural enemies can in some cases disrupt prey suppression, particularly when natural enemies engage in intraguild predation, where natural enemies compete with and prey upon each other. However, empirical studies have often demonstrated enhanced prey suppression despite intraguild predation. A recent theoretical study proposed the hypothesis that, when the intermediate predator is cannibalistic, intraguild predation can reduce cannibalism within the intermediate predator population, leading to little change in intermediate predator mortality and thus enhanced prey suppression. The goal of this study was to examine this hypothesis empirically. Two summer-long field enclosure experiments were conducted in cotton fields. We investigated the effects of adding an intraguild predator, Zelus renardii, on (1) the abundance of a cannibalistic intermediate predator, Geocoris pallens, (2) the abundance of a herbivore, Lygus hesperus, and (3) cotton plant performance. G. pallens adult abundance did not increase, even when food availability was high and natural enemies were absent, suggesting that density-dependent cannibalism imposes an upper limit on its densities. Furthermore, although Z. renardii is an intraguild predator of G. pallens, G. pallens long-term densities were unaffected by Z. renardii. In the presence of the intermediate predator, the addition of the intraguild predator Z. renardii enhanced suppression of L. hesperus, and there were suggestions that Z. renardii and G. pallens partitioned the L. hesperus population. Effects of herbivore suppression cascaded to the plant level, improving plant performance. In conclusion, we provide empirical support for the hypothesis that the addition of an intraguild predator may enhance prey suppression if the intermediate predator expresses density-dependent cannibalism. Intraguild predation and cannibalism co-occur in many communities; thus their joint effects may be broadly important in shaping predator effects on herbivores and plant performance.     

Re-creating missing population baselines for Pacific reef sharks

Sharks and other large predators are scarce on most coral reefs, but studies of their historical ecology provide qualitative evidence that predators were once numerous in these ecosystems. Quantifying density of sharks in the absence of humans (baseline) is, however, hindered by a paucity of pertinent time-series data. Recently researchers have used underwater visual surveys, primarily of limited spatial extent or nonstandard design, to infer negative associations between reef shark abundance and human populations. We analyzed data from 1607 towed-diver surveys (>1 ha transects surveyed by observers towed behind a boat) conducted at 46 reefs in the central-western Pacific Ocean, reefs that included some of the world's most pristine coral reefs. Estimates of shark density from towed-diver surveys were substantially lower (<10%) than published estimates from surveys along small transects (<0.02 ha), which is not consistent with inverted biomass pyramids (predator biomass greater than prey biomass) reported by other researchers for pristine reefs. We examined the relation between the density of reef sharks observed in towed-diver surveys and human population in models that accounted for the influence of oceanic primary productivity, sea surface temperature, reef area, and reef physical complexity. We used these models to estimate the density of sharks in the absence of humans. Densities of gray reef sharks (Carcharhinus amblyrhynchos), whitetip reef sharks (Triaenodon obesus), and the group "all reef sharks" increased substantially as human population decreased and as primary productivity and minimum sea surface temperature (or reef area, which was highly correlated with temperature) increased. Simulated baseline densities of reef sharks under the absence of humans were 1.1-2.4/ha for the main Hawaiian Islands, 1.2-2.4/ha for inhabited islands of American Samoa, and 0.9-2.1/ha for inhabited islands in the Mariana Archipelago, which suggests that density of reef sharks has declined to 3-10% of baseline levels in these areas.     

Predator and prey space use: dragonflies and tadpoles in an interactive game

Predator and prey spatial distributions have important population and community level consequences. However, little is known either theoretically or empirically about behavioral mechanisms that underlie the spatial patterns that emerge when predators and prey freely interact. We examined the joint space use and behavioral rules governing movement of freely interacting groups of odonate (dragonfly) predators and two size classes of anuran (tadpole) prey in arenas containing two patches with different levels of the prey's resource. Predator and prey movement and space use was quantified both when they were apart and together. When apart from predators, large tadpoles strongly preferred the high resource patch. When apart from prey, dragonflies weakly preferred the high resource patch. When together, large prey shifted to a uniform distribution, while predators strongly preferred the high resource patch. These patterns qualitatively fit the predictions of several three trophic level, ideal free distribution models. In contrast, the space use of small prey and predators did not deviate from uniform. Three measures of joint space use (spatial correlations, overlap, and co-occurrence) concurred in suggesting that prey avoidance of predators was more important than predator attraction to prey in determining overall spatial patterns. To gain additional insight into behavioral mechanisms, we used a model selection approach to identify behavioral movement rules that can potentially explain the observed, emergent patterns of space use. Prey were more likely to leave patches with more predators and more conspecific competitors; resources had relatively weak effects on prey movements. In contrast, predators were more likely to leave patches with low resources (that they do not consume) and more competing predators; prey had relatively little effect on predator movements. These results highlight the importance of investigating freely interacting predators and prey, the potential for simple game theory models to predict joint spatial distributions, and the utility of using model choice methods to identify potential key factors that govern movement.     

Influence of habitat on abundance and size structure of a large temperate-reef fish, Achoerodus viridis (Pisces: Labridae)

The relationship between densities of Achoerodus viridis (Pisces: Labridae) and reef habitats at various localities within New South Wales (NSW), Australia was examined. Types of habitats were quantified at inner, mid and outer estuarine locations in each of two estuaries (Botany Bay and Port Jackson) to determine whether purported patterns of movement from estuaries could be related to differences in habitat. Although the same types of habitat were generally found at all locations, differences in the proportion of habitat types were found between shallow and deep reefs and among inner, mid and outer estuarine locations for both estuaries. Shallow habitats were usually dominated by Ecklonia radiata, turf and/or fringe habitat in Botany Bay, whereas deep sites were generally dominated by urchin-grazed barrens habitat and, sometimes, sponge- and ascidian-dominated deep reef. Shallow sites in Port Jackson were dominated by a mixture of habitats, as were deep reefs at mid-estuarine locations. Other deep reefs in Port Jackson were dominated by E. radiata (inner estuarine) or barrens (outer estuarine) habitat. Thus, patterns of habitat cover were not consistent between estuaries and numbers of fish could not be related to proportional representation of habitat on reefs along estuarine gradients. Univariate and multivariate analyses showed that there was little evidence that any size class of fish was correlated with the proportional representation of a particular habitat or group of habitats. Counts of fish that focused on barrens and E. radiata forest habitats over a period of 10 yr showed that similar numbers and all sizes of fish were found in the two types of habitat. Greater numbers of small fish were, however, found in the E. radiata forest habitat than in the barrens habitat. Estimates of abundance along the coast of NSW (100s to 1000 km) in a range of habitats (e.g. ascidian-dominated reef, kelp forest, urchin-grazed barrens) showed that there was no indication that a particular habitat consistently had greater numbers of A. viridis than other habitats. Therefore, A. viridis of a range of sizes appears to be flexible in its use of habitats on reefs. Received: 24 December 1997 / Accepted: 23 June 1998     

Feeding ecology and evolutionary survival of the living coelacanth Latimeria chalumnae

One concept of evolutionary ecology holds that a living fossil is the result of past evolutionary events, and is adapted to recent selective forces only if they are similar to the selective forces in the past. We describe the present environment of the living coelacanth Latimeria chalumnae Smith, 1939 at Grande Comore, western Indian Ocean and report depth-dependent cave distribution, temperature, salinity and oxygen values which are compared to the fish's distribution and its physiological demands. We studied the activity pattern, feeding behaviour, prey abundance and hunting success to evaluate possible links between environmental conditions, feeding ecology and evolutionary success of this ancient fish. Transmitter tracking experiments indicate nocturnal activity of the piscivorous predator which hunts between approximately 200 m below the surface to 500 m depth. Fish and prey density were measured between 200 and 400 m, both increase with depth. Feeding tracks and feeding strikes of the coelacanth at various depths were simulated with the help of video and laser techniques. Along a 9447 m video transect a total of 31 potential feeding strikes occurred. Assuming 100% hunting success, medium-sized individuals would obtain 122 g and large females 299 g of prey. Estimates of metabolic rates revealed for females 3.7 ml O₂ kg⁻¹ h⁻¹ and for males 4.5 ml O₂ kg⁻¹ h⁻¹. Today coelacanths are considered to be a specialist deep-water form and to inhabit, with their ancient morphology, a contemporary environment where they compete with advanced, modern fish. Received: 5 July 1999 / Accepted: 11 November 1999     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

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.     

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.