Differential effects of habitat complexity,predators and competitors on abundance of juvenile and adult coral reef fishes

Greater structural complexity is often associated with greater abundance and diversity, perhaps because high complexity habitats reduce predation and competition. Using 16 spatially isolated live-coral reefs in the Bahamas, I examined how abundance of juvenile (recruit) and adult (non-recruit) fishes was affected by two factors: (1) structural habitat complexity and (2) the presence of predators and interference competitors. Manipulating the abundance of low and high complexity corals created two levels of habitat complexity, which was cross-factored with the presence or absence of resident predators (sea basses and moray eels) plus interference competitors (territorial damselfishes). Over 60 days, predators and competitors greatly reduced recruit abundance regardless of habitat complexity, but did not affect adult abundance. In contrast, increased habitat complexity had a strong positive effect on adult abundance and a weak positive effect on recruit abundance. Differential responses of recruits and adults may be related to the differential effects of habitat complexity on their primary predators. Sedentary recruits are likely most preyed upon by small resident predators that ambush prey, while larger adult fishes that forage widely and use reefs primarily for shelter are likely most preyed upon by large transient predators that chase prey. Increased habitat complexity may have inhibited foraging by transient predators but not resident predators. Results demonstrate the importance of habitat complexity to community dynamics, which is of concern given the accelerated degradation of habitats worldwide.     

Temporal variation in the diet of a coral reef piscivore (Pisces: Serranidae) was not seasonal

Seasonal and smaller scale temporal variations in the diet of the large predatory grouper Plectropomus leopardus were examined on the central Great Barrier Reef, Australia. In 5 months over a 2-year period, 742 P. leopardus (₅ cm standard length) were collected for stomach content analysis. Despite intensive sampling, identified prey were spread over 20 families and much of the dietary variation was considered to be stochastic. Most aspects of the diet of P. leopardus did not vary seasonally. Frequency, rates of feeding, and proportion of fish consumed were consistent throughout the study. Size of fish prey did not vary significantly between seasons. Pomacentridae, Caesionidae, Scaridae, and Labridae were four of the five most important prey families consumed and did not vary seasonally in the diet. Prey in the other family, the small-sized schooling Clupeidae, were consumed sporadically among months with no seasonal patterns. One less common prey family variedseasonally in the diet; Synodontidae, which are small voracious predators of fish recruits, were consumed by P. leopardus more in summer than winter. Although the diet of P. leopardus is not influenced by the seasonal influx of small fishes recruiting to reefs, these large piscivores may indirectly improve recruit survival by consuming small and abundant predators of new recruits.     

Food and feeding ecology of piscivorous fishes at Lake St Lucia, Zululand

The food and feeding ecology of piscivorous fish in Lake St Lucia was monitored for two years. Piscivorous fishes feed predominantly on the planktivorous Gilchristella aestuarius and Thryssa vitrirostris but a wide range of prey species was recorded. Numbers of the predominant piscivores, Argyrosomus hololepidotus and Elops machnata , in an area appear to be related to the densities of their major prey, T. vitrirostris and G. aestuarius . Large piscivorous fishes are restricted to the deeper portions of the lake, whereas small piscivores such as Johnius belengerii and Terapon jarbua feed predominantly on small fishes in the littoral zone. The highly significant correlation between the composition of prey fish species in the lake and prey fish species in the diet of piscivorous fishes, indicates that piscivores are feeding in a density dependent manner. However, factors such as habitat, fish size and swimming speed of prey species are shown to be important in prey selection. Juvenile fish of species such as Sarotherodon mossambicus, Liza macrolepis and Acanthopagrus berda remain in shallow marginal areas, thus avoiding large piscivorous fishes. However by frequenting shallow areas these species become vulnerable to bird predators, especially egrets and herons.     

Positive indirect effects in a coral reef fish community

Indirect effects occur when two species interact through one or more intermediate species. Theoretical studies indicate that indirect interactions between two prey types that share common predators can be positive, neutral or negative. We document a positive indirect interaction between different types of prey fish on coral reefs in Australia. A high abundance of one type of prey fish (cardinalfishes: Apogonidae) resulted in higher recruitment, abundance and species richness of other prey fish. Our evidence indicates that these effects were not due to differential settlement but were instead due to differential post-settlement predation. We hypothesize that resident piscivores altered their foraging behaviour by concentrating on highly abundant cardinal-fish when they were present, leaving recruits of other species relatively unmolested. Indirect effects were evident within 48 h of settlement and persisted throughout the 42-day experiment, highlighting the importance of early post-settlement processes in these communities.     

Prey selectivity of common predators on Silver carp (<Emphasis Type="Italic">Hypophthalmichthys molitrix</Emphasis>): controlled laboratory experiments support field observations

Silver Carp Hypophthalmicthys molitrix (Valenciennes, 1844) are an invasive species in the Mississippi River Basin and their current role in this novel ecosystem is not fully understood. Juvenile Silver Carp can and do occur in great numbers after a successful spawn. These massive schools of small Silver Carp seem to be an obvious prey source for the common predators of the Mississippi River system. The level to which native piscivores are consuming this novel prey item is unclear. Therefore, the goal of this research was to assess the diets of native piscivores collected in Pool 26 and the Open River reach of the Mississippi River. Using diet contents and catch rates of small fishes, selection or avoidance could be determined for predator prey interactions. Then a controlled laboratory experiment was conducted to determine if common predators [White Bass Morone chrysops and Largemouth Bass Micropterus salmoides (Lacepede, 1802)] would select for or against this novel prey in the presence of two native prey fish [Gizzard Shad Dorosoma cepedianum (Lesueur, 1818) and Emerald Shiner Notropis atherinoides]. Understanding how predator-prey interactions occur in a controlled laboratory experiment may provide insight to trends observed in the field. The majority of native piscivores appeared to be avoiding Silver Carp in both reaches and selecting native prey fishes. The order in which prey fish were consumed by both predators showed significant avoidance of Silver Carp. It appears that the trends observed in the field were supported by our controlled laboratory experiment. The effectiveness of different native piscivores to consume Silver Carp may have impacts on future management decisions.     

Consumers of sea urchins,<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> and<Emphasis Type="Italic"> Arbacia lixula</Emphasis>, in shallow Mediterranean rocky reefs

Underwater observations on fish and asteroid consumers (i.e. predators and scavengers) of sea urchins, Paracentrotus lividus and Arbacia lixula, were carried out at several locations in shallow Mediterranean rocky reefs. Observations conducted in the marine reserve of Torre Guaceto (Adriatic Sea) revealed that sparid fishes, Diplodus sargus and D. vulgaris, are the main fish predators of small (<1 cm in test diameter) and medium (1–4 cm) sea urchins, whereas the labrids Coris julis and Thalassoma pavo preyed only upon small sea urchins. Large D. sargus were able to prey upon small and medium, and occasionally large (>4 cm) sea urchins, whereas medium and small Diplodus preyed mainly upon small sea urchins. The number of sea urchins preyed upon by fishes was negatively related to sea urchin size for both species. P. lividus appeared to be subject to higher predation levels than A. lixula. The scavenger guild comprised 11 fish species, with D. sargus, D. vulgaris, Coris julis and Chromis chromis accounting for about 80% of scavenger fishes. Observations performed at several locations in the Mediterranean on the predatory asteroid Marthasterias glacialis revealed that only 3% of the detected individuals were preying upon sea urchins. Due to the importance of sea urchins for assemblage structure and functioning of Mediterranean rocky reef ecosystems, these results may have also important implications for management of fishing activities.Communicated by H.-D. Franke     

Fish larvae distribution among different habitats in coastal East Africa

Fish larvae abundances, diversity and trophic position across shallow seagrass, coral reef and open water habitats were examined to characterize their distribution in coastal East Africa. Larvae were identified to family and analysed for abundance differences between sites and habitats, trophic level using stable-isotope analysis and parental spawning mode. Abundances differed greatly between sites with the highest numbers of larvae occurring in the open-water and seagrass habitats. Larval fish diversity was high across habitats with 51 families identified with small differences between sites and among habitats. Notably, larvae of abundant large herbivorous fishes present in reef and seagrass habitats were almost completely absent at all sampling locations. In the seagrass, demersal spawned larvae were more abundant compared with the reef and open-water habitats. Stable-isotope analysis revealed that fish larvae have a varied diet, occupying trophic level two to three and utilizing planktonic prey. This study offers new insights into distributional aspects of fish larvae along the East African coast where such information is sparse.     

Diet niche relationships among predator and prey fish species in their early life stages in Lake Võrtsjärv (Estonia)

In Lake Võrtsjärv pikeperch was observed not to shift to piscivory in their first autumn of life, although juvenile stages of a variety of fish species were abundant in the lake. It was hypothesized that the diets of predator and prey fish fry overlap and that coarse fish species are important food competitors for juvenile piscivores and thus, pikeperch and perch fry do not shift to piscivory during their first growing season. To discover the possible linkages in this pattern, in 2009 the feeding relationships of pikeperch, perch, ruffe and roach fry were analysed. The stomach content analyses showed that in the summer period, Mesocyclops leuckarti was the most frequent prey for perch and ruffe, pikeperch consumed Leptodora kindti in large quantities, and roach ate mostly plant material. Towards autumn, M. leuckarti was the most abundant prey for all percids. However, average stomach content weight and the number of prey items eaten by ruffe were considerably higher than for other fish fry. Since the feeding opportunities of fish fry are considered poor in the examined lake, the prey has the potential to restrict the recruitment to piscivory of their predators, as prey fish seem to have better abilities to persist in this ecosystem. Furthermore, supposed competition in the juvenile stage may result in a reduced top‐down effect on coarse fish.     

Ontogenetic changes in foraging tactics of the piscivorous cornetfish <Emphasis Type="Italic">Fistularia commersonii</Emphasis>

Foraging behaviors of the piscivorous cornetfish Fistularia commersonii were observed at shallow reefs in Kuchierabu-jima Island, southern Japan. This fish foraged on two types of prey fishes: one was reef fish that typically dwell on or near substrata (e.g., Tripterygiidae and Labridae), and the other was pelagic fish that shoal in the water column (e.g., Clupeidae and Carangidae). The prey sizes, prey types and foraging behaviors changed as the predator size increased. Prey sizes were largely limited by gape size of the cornetfish, and small predators consumed small prey. The small cornetfish (10–30 cm in total length) fed only on reef fish captured after stalking (where the fish slowly approaches the prey and then suddenly attacks). The stalking was done either solitarily or in foraging association with conspecifics. Large fish (30–120 cm) fed on both types of fishes by stalking and/or chasing (where the fish chases the prey using its high mobility and attacks), either solitarily or in foraging association with con- or heterospecifics. Thus, chasing was only performed by the large cornetfish against pelagic prey fish in associative foraging with other con- and heterospecific predators. As their body sizes increased, F. commersonii began to show a diversification of foraging behaviors, which was strongly related not only to the habitat types and anti-predatory behaviors of the prey fishes but also to associative foraging with con- or heterospecifics, which improves their foraging success.     

Temporal patterns of resource partitioning among Cichla species in a Venezuelan blackwater river

In channel and floodplain habitats of the Cinaruco River, Venezuela, Cichla temensis was more abundant and larger than C. intermedia and C. orinocensis . Seasonal variation in hydrology influenced habitat use, spawning, and predator-prey interactions. The three piscivores partitioned habitat, with C. intermedia showing a strong affinity for structured habitats in the main channel during all water level fluctuations. C. orinocensis was most abundant in shallow areas with submerged structure in lagoons and, to a lesser extent, in low velocity regions of the channel, and C. temensis occupied a wide range of lotic and lentic habitats. During the low-water period, the feeding frequency and body condition of all three species declined, and this was related, in part, to preparation for spawning near the end of the low-water season. The diet of C. intermedia was least similar to its two congeners during falling and rising water. C. orinocensis and C. temensis had lowest diet overlap during the low-water conditions, the period when many individuals of these two species move into lagoons for nesting. Prey in stomachs were significantly larger during the falling-water than the rising-water period, and predation by Cichla and other large piscivores during the falling-water period may have reduced the abundance of large prey, particularly Semaprochilodus kneri . These migratory detritivorous fish were important prey for C. temensis during the falling-water period and probably contributed a substantial fraction of the annual energy intake for this species. Together, the three Cichla species consume a wide spectrum of prey from a diverse fish assemblage, but prey are subdivided based on habitat, prey type, and season.     

Dietary shifts in a stressed fish assemblage: Consequences of a bivalve invasion in the San Francisco Estuary

We compared dietary patterns within a temperate estuarine fish assemblage (Suisun Marsh, CA, U.S.A.) during a period of high mysid shrimp abundance and after a major decline in mysid abundance caused by the invasion of the overbite clam Potamocorbula amurensis. Prior to the invasion, high dietary overlap, high stomach fullness, and low niche breadth occurred among the fishes in spring when mysid populations were high. Dietary overlaps decreased and niche breadth increased for all species but the endemic splittail Pogonichthys macrolepidotus in fall when mysid populations were low. Eight native species exhibited lower overall collective overlaps and fuller stomachs than five alien species, suggesting more efficient resource partitioning. After mysid abundance declined, only alien striped bass Morone saxatilis preyed upon mysids in greater than trace amounts. An alien mysid became an important prey for small striped bass, but striped bass also switched to piscivory at a smaller size than when mysids were abundant. Eight of 13 species exhibited significant declines in abundance during the study period, which were concordant with the original importance of mysids in their diets. Our results suggest that altered lower food web dynamics in the San Francisco Estuary caused by the invasion of the overbite clam changed fish diets and have contributed to declines in fish abundance.     

Interactive effects of habitat selection, food supply and predation on recruitment of an estuarine fish

 Seagrass meadows are often important habitats for newly recruited juvenile fishes. Although substantial effort has gone into documenting patterns of association of fishes with attributes of seagrass beds, experimental investigations of why fish use seagrass habitats are rare. We performed two short-term manipulative field experiments to test (1) the effects of food supply on growth and densities of fish, and (2) effects of predation on the density and size distribution of fish recruits, and how this varies among habitat types. Experiments were conducted in Galveston Bay, Texas, and we focused on the common estuarine fish, pinfish Lagodon rhomboides. In the first experiment, replicate artifical seagrass and sand plots were either supplemented with food or left as controls. Recruitment of pinfish was significantly greater to seagrass than sand habitats; however, we detected no effect of food supplementation on the abundance of recruits in either habitat. Pinfish recruits in artifical seagrass grew at a significantly faster rate than those in sand habitats, and fish supplemented with food exhibited a greater growth rate than controls in both sand and artifical grass habitats. In our second experiment, we provided artificial seagrass and sand habitats with and without predator access. Predator access was manipulated with cages, and two-sided cages served as controls. Recruitment was significantly greater to the cage versus cage-control treatment, and this effect did not vary between habitats. In addition, the standard length of pinfish recruits was significantly larger in the predator access than in the predator exclusion treatment, suggesting size-selective predation on smaller settlers or density-dependent growth. Our results indicate that the impact of predation on pinfish recruits is equivalent in both sand and vegetated habitats, and thus differential predation does not explain the higher recruitment of pinfish to vegetated than to nonvegetated habitats. Since predators may disproportionately affect smaller fish, and a limited food resource appears to be more effectively utilized by fish in vegetated than in unvegetated habitats, we hypothesize that pinfish recruits may select vegetated habitats because high growth rates allow them to achieve a size that is relatively safe from predation more quickly. Received: 10 October 1996 / Accepted: 5 April 1997     

Habitat-related predation on juvenile wild-caught and hatchery-reared red drum Sciaenops ocellatus (Linnaeus)

We examined the patterns of habitat-specific mortality for newly settled red drum (Sciaenops ocellatus) using an experimental mesocosm approach. Experiments were designed to analyze prey vulnerability and fish rearing-type (wild-caught or hatchery-reared) in estuarine habitats of varying structural complexity including marsh (Spartina alterniflora Loisel), oyster reef (Crassostrea virginica Gmelin), seagrass (Halodule wrightii Aschers), and nonvegetated sand bottom. We used two different predators, pinfish (Lagodon rhomboides Linnaeus) and spotted seatrout (Cynoscion nebulosus Cuvier). For both predators, vulnerability of wild-caught red drum was significantly lower in structurally complex habitats such as seagrass and oyster reef; the highest vulnerability was associated with the nonvegetated bottom. This habitat effect was not apparent for hatchery-reared prey. In trials using a combination of both rearing-types, there was no significant habitat effect on prey selection, but hatchery-reared red drum suffered higher overall mortality than wild-caught fish from pinfish predators. In these trials, spotted seatrout did not select for either prey type. Differences we observed in prey vulnerability were likely caused by behavioral differences between wild-caught and hatchery-reared red drum. Our results reinforce the conclusion that structural complexity in estuarine habitats increases survival of newly settled fishes. Our data also suggest that hatchery-reared red drum may be more vulnerable to predation than natural fishes, and that survival of stocked fish may be enhanced through habitat-related behavior modification.     

Patterns of abundance variation in reef fishes near an artificial reef at Guam

Synopsis Development of the fish community on a submerged 16 m barge and variation in fish abundance on nearby transects were surveyed twice monthly for twenty months. A steady increase in abundance was observed for certain fishes on the barge, whereas a few species exhibited distinct seasonal variation on both the barge and transects. Most of the seasonal species settled between March and May.Some seasonal species appeared to be site selective in their settlement and consequently settled juveniles were clumped in their distribution. An abundance of preferred topographical features may be why settlement was relatively high at the study site and indirectly why predators became significantly (r₃ = 7.67^***, N = 37) more abundant at the study area during the months of maximum prey settlement. Concurrent settlement of several species during the same few months may be important because juveniles become an abundant food source to predators during those few months only. Periodic swamping of predators by abundant juvenile prey may improve the chances for individuals of rarer prey species to be overlooked and therefore be succesfully recruited.     

Vulnerability of marine forage fishes to piscivory: effects of prey behavior on susceptibility to attack and capture

We conducted a series of size-structured laboratory experiments to quantify and compare the susceptibility of several estuarine and marine forage fishes to attack and capture by piscivorous predators. Size-dependent estimates of capture success, handling time, and prey profitability were generated from single-species experiments offering bay anchovy, Atlantic menhaden, Atlantic silverside, and age-0 striped bass to piscivores. Bay anchovy and Atlantic menhaden were most susceptible to capture and yielded high profitability compared to Atlantic silverside and age-0 striped bass prey. Variation in capture success among forage species was particularly influential in generating disparate profitability functions. Although morphological differences among forage species contributed to variation in susceptibility to predation, behavioral analyses indicated that variable reaction distances to approaching predators and activity levels of prey may explain a large fraction of the observed differences in susceptibility. When several forage species were offered to predators simultaneously in larger enclosures, mortality was highest and occurred earlier for bay anchovy and Atlantic menhaden compared to other prey, which points to the strong influence of predator capture success on overall forage fish vulnerability. Our results demonstrate species-specific differences among forage fishes in the ability to avoid attack and capture by piscivores, and we conclude that the expression of antipredator behaviors contributes significantly to variation in forage species vulnerability.     

Is there safety‐in‐numbers for prey?

The abundance of prey affects the rate of predation, but little consensus exists on whether this enhances or reduces per capita mortality. Studies of aggregating prey in marine habitats generally emphasise that the probability of predation of any individual is the reciprocal of the number of prey within a school. A field experiment tested the alternative hypotheses that predation by predatory fish on schooling prey (1) increased with an increase in the number of prey per school and that this caused (2) survival to be lower in schools with more individuals. The number of prey (juvenile Acanthochromis polyacanthus ) per school was manipulated in replicate treatments with natural densities of large predatory fish (open plots) and treatments without large predatory fish (exclusion cages). Large predatory fish preyed on juveniles in a density-dependent manner and this was the key source of density-dependent mortality in plots open to all predators. There was some suggestion that small predatory fish also prey on juveniles in a density-dependent manner, but this was weak and did not translate into density-dependent mortality of juveniles. It would appear that aggregation of prey may be a successful strategy against predation from some predators, but not always every predator, or all predators in combination.     

The influence of prey communities on fish species assemblages on artificial reefs in Puget Sound,Washington

Synopsis Research on eleven artificial reefs in Puget Sound, Washington examined the relative importance of reef-produced prey items to recreationally important reef fish species assemblages. The colonization of potential prey items, and fish species assemblages to ten artificial reefs were examined for the reefs first two to five years, and observations were conducted on an eleventh reef during its forty-ninth productive year. Fish species became more abundant, or were seen more frequently on reef habitats whose substrates had successionally developed from barnalces to algal mats. Fish species most affected by this successional change foraged heavily on organisms which were associated with reef algae. Starfish and nudibranchs. who preyed on the barnacles, were identified as the ‘keystone’ predators of these subtidal reef habitats.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua , hake Merluccius merluccius , megrim Lepidorhombus whiffiagonis , whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey ( e.g . Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey ( e.g . mackerel Scomber scombrus ), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes ( Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

Small cryptopredators contribute to high predation rates on coral reefs

Small fishes suffer high mortality rates on coral reefs, primarily due to predation. Although studies have identified the predators of early post-settlement fishes, the predators of small cryptobenthic fishes remain largely unknown. We therefore used a series of mesocosm experiments with natural habitat and cryptobenthic fish communities to identify the impacts of a range of small potential predators, including several invertebrates, on prey fish populations. While there was high variability in predation rates, many members of the cryptobenthic fish community act as facultative cryptopredators, being prey when small and piscivores when larger. Surprisingly, we also found that smashing mantis shrimps may be important fish predators. Our results highlight the diversity of the predatory community on coral reefs and identify previously unknown trophic links in these complex ecosystems.

     

Shallow water predation risk for a juvenile flatfish (winter flounder; Pseudopleuronectes americanus, Walbaum) in a northwest Atlantic estuary

Many small fish, including several juvenile Atlantic flatfish, are most abundant in shallow areas presumable because these habitats enhance survivorship and/or growth. In this study, we investigated size-dependent depth distributions and the role of shallow habitats as predator refuges for age-0 winter flounder (Pseudopleuronectes americanus) in a northwest Atlantic estuarine nursery. Analysis of trawl surveys performed during the larval settlement period throughout the Navesink River and Sandy Hook Bay, New Jersey, showed that as fish increased in size, depth of occurrence gradually decreased, so that individuals >35 mm standard length (SL) were concentrated in habitats ∼1 m deep. Tethering in structurally simple and adjacent shallow and deep habitats showed that predation risk for flounder (30-50 mm SL) was low in shallow water (<1 m) and increased rapidly with depth. Summer flounder (Paralychthys dentatus), which were more abundant in trammel nets in deep habitats and included winter flounder in their diets, appeared to be important consumers of tethered fish. Our results indicate that following larval settlement, winter flounder emigrate from or suffer high mortality in deeper water to become concentrated in shallow habitats that can serve as predator refuges even when they lack complex physical structures. These results highlight the potential for functional habitat loss when natural and/or anthropogenic factors make shallow habitats unavailable to young fish.