Effects of frequent fish predation on corals in Hawaii

The abundance of lesions from fish bites on corals was quantified at nine shallow reefs in the main Hawaiian Islands. There were on average 117 bite scars m⁻² on Pocillopora meandrina tissue from the barred filefish Cantherhines dumerilii, 69 bites m⁻² on Porites compressa tissue, and 4 bites m⁻² on Porites lobata tissue from the spotted puffer Arothron meleagris. Across sites, the frequency of A. meleagris bites on P. compressa per unit area of living coral cover declined exponentially with increasing coral cover. P. compressa nubbins in two size classes (1–2 cm and 4–5 cm) were transplanted onto six study reefs. Nubbins in the small size class were entirely removed by bites from A. meleagris, while nubbins ≥4 cm were only partially consumed, leaving them able to recover. At sites with abundant P. compressa, predation had little effect on transplanted nubbins; at sites where P. compressa comprised less than 5% of living cover, all nubbins were preyed upon. A. meleagris bite lesions on P. compressa were monitored through time and fully recovered in 42 ± 4 days. A model of the risk of over-predation (a second predation event before the first is healed) decreased exponentially with increasing coral cover and increased linearly with increasing lesion healing time. The increased risk of over-predation at low coral cover could indicate an Allee effect limiting the recovery of coral populations if coral cover is substantially reduced by natural or anthropogenic disturbances.     

The relative importance of predation and competition in two reef fishes

Competition and predation may both strongly influence populations of reef fishes, but the importance of these processes relative to one another is poorly understood. I quantified the effects of predation and competition on the growth and survival of two temperate reef fishes, Lythrypnus dalli and Coryphopterus nicholsii, in field experiments in which I manipulated the densities of the two species and the abundance of predators (using exclosure cages) on small replicate patch reefs. I also evaluated the influence of predators on the behavior of the two species to help interpret the mechanisms of any predatory influences on growth or survival. Predation was much more important than competition (inter- or intraspecific) in Lythrypnus. For Coryphopterus, neither competition nor predation were particularly important. Behaviorally, both species responded to predators by reducing foraging rate and hiding. This altered behavior, however, had no repercussions for growth or survival of Coryphopterus. In contrast, Lythrypnus grew more slowly and suffered greater mortality when exposed to predators. Interspecific competition did not significantly influence either species. Intraspecific competition did not affect the growth of Coryphopterus, but survival tended to be lower at high densities. Growth of Lythrypnus was depressed by intraspecific competition, but survival was not, except that, in the presence of predators, survival was density dependent. In contrast to the historical emphasis placed on the role of competition, this study indicates that predation can be more important than competition in determining patterns of abundance of some reef fishes. For example, predators not only influenced foraging of both Lythrypnus and Coryphopterus, but they also reduced growth and survival of Lythrypnus, and therefore appear to help maintain the marked habitat segregation between the two species. Received: 16 June 1997 / Accepted: 3 December 1997     

Competition between scleractinian corals and macroalgae: An experimental investigation of coral growth, survival and reproduction

Macroalgae are a major component of many coral reef flat communities, and are potentially major competitors with corals. The influence of macroalgae on several demographic parameters of four species of scleractinian coral by means of an algal clearance experiment was examined to determine specifically if macroalgae are affecting coral cover, growth, fecundity, fission, survivorship and recruitment. Also investigated were patterns of natural encounters between corals and algae.

Algal cover at the study site ranged from 41 to 56%, and coral cover from 8 to 10%. In total, 92 ± 4 ( )% of coral colonies were in contact with one or more species of macroalgae. Changes in coral cover were significantly affected by the presence of macroalgae, with cover of Acropora species increasing faster in areas from which algae had been cleared compared to control areas where algae had not been removed, although this pattern did not occur for Pocillopora damicornis (Linnaeus). Similarly, growth of individual colonies was faster when macroalgae were absent for three Acropora species but not for P. damicornis. There were no differences detected in rates of fission or survivorship of corals between algal clearance and control treatments, although there were high levels of variability in both of these parameters. Fecundity of Acropora palifera (Lamarck), the only species examined, was approximately double in colonies in cleared plots compared to those in control plots with macroalgae present. As no recruitment occurred throughout the 2-yr study, it remains to be determined how macroalgae effect the settlement of coral larvae. The results show that macroalgae can have a major influence on the demography of scleractinian corals.     

The influence of habitat characteristics and conspecifics on attraction and survival of coral reef fish juveniles

In marine species with a pelagic larval stage, search behavior and selection of a suitable reef habitat can maximize the settlement success of recently settled juveniles and their subsequent performance (growth and survival of juveniles). Our objective was to test this hypothesis for a single target coral reef fish species (Chromis viridis) at Moorea Island. C. viridis settle on living coral colonies of Porites rus already populated with conspecifics. In the present study (conducted in experimental cages), we found that: 1) mortality rate of recently settled juveniles of C. viridis was lower in the settlement habitat (living coral colonies of P. rus) than in other habitats having physical structure different from those of P. rus colonies; 2) C. viridis juveniles preferentially colonized coral heads of P. rus with conspecifics present rather than uninhabited coral heads and they also preferentially colonized uninhabited coral heads rather than coral heads with heterospecifics; 3) mortality rate of C. viridis juveniles did not vary with the presence or absence of conspecifics or heterospecifics on P. rus colonies. Overall, the study allows us to highlight that site selection by juveniles for habitat containing conspecifics does not benefit their short term mortality rates, suggesting that in the short term at least, site selection has little importance.     

Effects of larval competition on survival and growth in Mediterranean fruit flies

1. Larval success was compared when one, two, or three egg clutches were laid in kumquat fruits (≈ 10 ml in volume) either successively on the same day or at the rate of one clutch per day. 2. Increased clutch density was associated with a significant decrease in larval survival rate and non‐significant decreases in larval growth rate and pupal mass. 3. Larval and pupal parameters showed significantly larger variance when clutches were laid on successive days than on the same day, suggesting a competitive advantage for older larvae over younger larvae. 4. The results suggest that, in small fruit, reduced fitness due to larval competition may act against possible fitness benefits due to social facilitation among adult females, hence reducing the likelihood of non‐linear population dynamics caused by processes such as the Allee effect.     

Relative importance of predation and intraspecific competition in regulating growth and survival of juveniles of the soft-shell clam, Mya arenaria L., at several spatial scales

Predation appears to be the single most important biotic factor regulating populations of bivalves in estuarine and coastal soft sediments. However, the relative roles of predation and intraspecific competition are rarely investigated simultaneously over different spatial scales, making generalities about these mechanisms difficult. Using juveniles of the soft-shell clam, Mya arenaria (initial mean shell length [SL] ± 95% CI = 12.4 ± 0.13 mm), I tested the interactive effects of predator deterrence and intraspecific density (660 vs. 1320 individuals m^− 2) on growth and survival responses over a 185-day period from May to November 2003 at spatial scales that spanned four orders of magnitude: embayments, sites within embayments, tidal gradients, and blocks that were 10,000's, 1000's, 100's, and 5 m apart, respectively. Replicate field experiments were conducted from May to November 2003 at the upper and lower tidal heights at each of two intertidal mud flats (sites) within each of two embayments (Passamaquoddy Bay [PB] and Cobscook Bay [CB]) in eastern Maine.Mean survival, relative growth, and the abundance of wild recruits each varied significantly over all spatial scales. Predation was the most important factor affecting clam survival, explaining 45% of the total variability, whereas embayment, sites within embayments, tidal gradient, and intraspecific density collectively accounted for less than 10% of the variation. At all four intertidal sites, clam survival in experimental units designed to deter predators averaged 72%, but the degree of enhancement varied between embayments (PB = 61%; CB = 267%). Average survival rate was higher (by 12-16%), but growth was slower (by ca. 50%) in upper vs. lower intertidal blocks; however, the patterns differed for both variables between sites within each embayment. The effect of increasing intraspecific clam density was to lower survival by ca. 17% (56% [660 m^− 2] vs. 48% [1320 m^− 2]) in both embayments, but growth was unaffected. Overall, clams doubled in SL, although mean relative growth was 15% greater in CB than PB. Tidal gradient, sites within embayments, and blocks were the three most important factors explaining 35%, 19%, and 22% of total variation in relative clam growth, respectively. In Maine and the northeast US, juveniles of Mya reach their highest abundance above mean low tide levels. Experimental evidence presented here suggests that differential predation along the tidal gradient is the dominant factor controlling clam abundance and distribution patterns in the intertidal zone.     

Effects of macroalgal exudates and oxygen deficiency on survival and behaviour of fish larvae

Filamentous algae may create anoxic, i.e. oxygen free, conditions during night and fish larvae that commonly spend their first months in the littoral may therefore be subject to anoxia. We conducted two different experiments; firstly, we measured behaviour of pike larvae (Esox lucius L.), such as frequency of prey attacks and time in vegetation, in the presence of a chemical predator cue and oxygen deficiency. In the second experimental set-up, pike larval survival was monitored in water with added macro-algal exudates (excreted from Pilayella littoralis) and a low oxygen level (3 mg l^− 1). Our results showed that oxygen concentration and the chemical predator signal, caused by three-spined sticklebacks, had strong significant effects on the larval prey attacks. The prey attacks were strongly reduced at 3 mg oxygen l^− 1. However, survival of the larvae was not affected by low oxygen and macro-algal exudates. Pike larvae are very tolerant to hypoxia and exudates excreted by the littoral vegetation, whereas prey attacks are strongly decreased in hypoxia when a predator is around. This may have negative consequences for pike larval growth and recruitment in eutrophicated environments.     

Trade-offs in the response of mayflies to low oxygen and fish predation

Summary We examined how mayfly larvae (Ephemeroptera,Callibaetis montanus) balance the conflicting demands of avoiding both benthic hypoxia and fish predators. Using vertical oxygen and temperature gradients typical of ice-covered lakes, we observed the behavior of mayflies in the presence and absence of fish. In the absence of fish and with adequate oxygen, mayflies spent most of the time on the bottom substrate. As benthic oxygen concentration declined, mayflies increased their activity and moved up in the water column. In the presence of fish and with adequate oxygen, mayflies spent even more time associated with the bottom substrate and reduced their activity levels. As benthic oxygen concentrations declined, mayflies increased their activity and moved up in the water column, but to a lesser extent than when fish were absent. Because of this depression in activity and reluctance to leave the bottom substrate, mayflies endured lower oxygen concentrations in the face of predation threat relative to when fish were absent. Despite this trade-off, benthic hypoxia resulted in increased mortality due to fish predation. Because benthic invertebrates vary in their ability to tolerate hypoxia and in their vulnerability to fish predators, periods of benthic hypoxia could lead to selective predation on some taxa and be an important force structuring benthic invertebrate assemblages.     

Regeneration of artificial injuries on scleractinian corals and coral/algal competition for newly formed substrate

Porites cylindrica and Porites lutea fragments of colonies were inflicted with five different injury types: chisel, file, Water Pik, osmotic and cement injuries. The fragments were maintained in outdoor aquaria for a period of 240 days under light intensities varying from 2-5% to 70-90% of incident surface photosynthetic active radiation (PAR₀). During the exposure, changes in weight of the fragments, the rates of regeneration of the injuries, abundance of algae and animals settled onto injured areas were monitored. The regeneration rate of the injuries depended on interspecific differences in corals, injury types, number and composition of algae and animals settled onto the lesions, and light and temperature conditions. Competitive interactions between polyps and settlers occurred after colonizers settled onto the damaged surface or the live tissue. It is noteworthy that recovered coral tissue generally overgrew about 100 algal species with or without inhibition of coral growth by algae. In the summer period, the cyanobacterium Lyngbya majuscula covered some lesions (osmotic and cement) by 100%, thus reducing dramatically the regeneration rate of the inflicted injuries and also caused coral bleaching when in direct contact.     

Transmission of growth anomalies between Indo-Pacific Porites corals

In this study we provide experimental evidence of transmission of growth anomalies (GAs) between corals. Twenty-four aquaria (16 experimental, 8 containing only apparently healthy corals) were set-up on Negros Island, Philippines, to test for direct-contact and waterborne transmission of GAs. Within seven weeks, two of 16 apparently healthy colonies placed in direct contact with colonies having GAs developed multiple GA lesions whose size and number increased over time. One of 16 apparently healthy colonies in experimental aquaria not touching any diseased colony also developed a GA, exhibiting a single lesion that did not increase in size. Apparently healthy colonies (n=24) in aquaria without a diseased colony remained unchanged.     

Habitat complexity modifies the impact of piscivores on a coral reef fish population

Patterns in juvenile mortality rates can have a profound affect on the distribution and abundance of adult individuals, and may be the result of a number of interacting factors. Field observations at Lizard Island (Great Barrier Reef, Australia) showed that for a coral reef damselfish, Pomacentrus moluccensis, juvenile mortality (over 1 year) varied between 20 and almost 100% among sites. Correlative data showed that juvenile mortality increased as a function of initial densities (recruitment), predator densities and the availability of preferred coral substrata. A multiple regression showed that these three variables together did not explain significantly more variation in mortality than the single factor showing the strongest relationship. This appeared to be because recruitment, predator densities and preferred coral substrata were all highly correlated, suggesting that one, two or all of these factors may be influencing juvenile mortality rates. One hypothesis was that density-dependent mortality in juveniles was the result of an interaction between predators (which appear to aggregate at high-recruitment sites) and the availability of preferred substrata (predator refuges). We tested this hypothesis by using both laboratory and field experiments to see whether fish predation could significantly alter survivorship of this damselfish, and whether this impact was dependent upon the coral substratum. The laboratory experiment was designed to test the effects of three common predators (Pseudochromis fuscus, Cephalopholis boenak and Thalassoma lunare) and three different coral substrata that varied in their complexity (Pocillopora damicornis, Acropora nasuta and A. nobilis) on the survival of juvenile Pomacentrus moluccensis. There was a significant interaction between predator species and microhabitat in determining survival. Pseudochromis fuscus and C. boenak were both significantly better at capturing juvenile damselfish than T. lunare. Juvenile survivorship was significantly better when they were given the more complex corals, Pocillopora damicornis and A. nasuta, compared with those given the open-structured species A. nobilis. This pattern reflects habitat selection in the field. Predators differed in their strike rates and the proportion of strikes that were successful, but all exhibited greater success at prey capture where A. nobilis was provided as shelter. The interaction between the effect of predator species and microhabitat structure on damselfish survival was tested in the field for a cohort of juvenile Pomacentrus moluccensis. We examined juvenile survival in the presence and absence of two predators that co-occur on natural patch reefs (C. boenak and Pseudochromis fuscus). The experimental patch reefs we used for this purpose were constructed from both high complexity (Pocillopora damicornis) and low complexity (A. nobilis) coral substrata. Both juveniles and predators were translocated to reefs at natural densities. The effects of predation were clearly dependent upon the microhabitat. Reefs of the high-complexity coral with predators supported the same high numbers of Pomacentrus moluccensis as the reefs with no resident predators. However, damselfish abundance was significantly lower on low-complexity reefs with resident predators, relative to the other treatments. Background rates of loss were high, even on preferred coral in the absence of the manipulated predator, suggesting that transient predators may be even more important than the residents. We suggest that adult abundances in this species were strongly influenced by the densities of different predators and the availability of preferred refuges. Received: 3 April 1997 / Accepted: 26 August 1997     

Scorpions,spiders and solpugids: predation and competition among distantly related taxa

Summary Scorpions, spiders and solpugids are generalist predators on the same types of arthropod prey. However, these potential competitors also frequently eat one another (=intraguild predation). In a 29 mo. experiment, >6,000 scorpions were removed from 300 (10x10m) quadrats. Significantly more spiders (but not solpugids) occurred in removal versus control quadrats. Two alternate hypotheses potentially explain this result: exploitation competition for jointly exploited prey or intraguild predation. There was no evidence of exploitation competition: no differences existed between removal and control quadrats in prey abundance or spider size and reproductive characteristics. It appears that the release from predation pressure in areas from which scorpions were removed produced the observed increase in spider abundance. Current ecological theory does not fully apply to situations whereby species at the same trophic level interact as both predators and potential competitors.     

Interactive effects of interspecific competition and microhabitat on early post-settlement survival in a coral reef fish

Microhabitat type and the competition for microhabitats can each influence patterns of abundance and mortality in coral reef fish communities; however, the effect of microhabitat on the intensity and outcome of competition is not well understood. In Kimbe Bay, Papua New Guinea, surveys were used to quantify microhabitat use and selectivity in two live-coral specialist damselfishes (Pomacentridae), Chrysiptera parasema, and Dascyllus melanurus. A patch reef experiment was then conducted to test how intra- and interspecific competition interacts with two types of microhabitat to influence survival of recently settled C. parasema. Surveys demonstrated that C. parasema and D. melanurus recruits utilized similar coral microhabitats; 72% of C. parasema and 85% of D. melanurus used corymbose and bottlebrush growth forms of Acropora. One microhabitat type, Pocillopora sp. coral, was commonly used by D. melanurus but rarely by C. parasema. The patch reef experiment revealed that both microhabitat and interspecific competition influence abundance of recently settled C. parasema. Microhabitat had the strongest influence on survival of C. parasema. In the absence of interspecific competitors, ~85% of C. parasema survived for 5 days after transplantation to high-complexity bottlebrush Acropora reefs when compared to only 25% survival of Pocillopora reefs. In both microhabitats, interspecific competition with D. melanurus, but not intraspecific competition, significantly decreased the survival of C. parasema. Taken together, these results suggest that the observed distribution of C. parasema results from specialized microhabitat requirements and competition for space in those microhabitats. This study demonstrates that interspecific competition and microhabitat type can interact to influence early post-settlement survival in coral reef fishes, though, whether and how these factors influence survival will depend on the behavioural attributes and strength of habitat associations among potential competitors. Communicated by Environment Editor Prof. Rob van Woesik     

The effects of predation by planktivorous juvenile fish on the microbial food web

The feeding impact of planktivorous fish on microbial organisms is still poorly understood. We followed the seasonal dynamics of the food web in two natural fishponds for two years: one was stocked with planktivorous whitefish while the other had no planktivorous fish. The aim of the study was the simultaneous assessment of the feeding behaviours of planktivorous fish and of bacterivorous meta-/protozooplankters. We hypothesized that in the presence of planktivorous fish there would be fewer metazooplankton, more protozoans and decreased numbers of bacteria. Our results showed that the amount of metazooplankton eaten by the fish was indeed negatively correlated with metazooplankton biomass. The feeding impact of planktivorous fish in shaping the microbial loop was remarkable. The main grazers of bacteria in the fishpond were ciliates, whereas in the pond without fish these were heterotrophic nanoflagellates. In the fishless pond the role of the top predator shifted to the predaceous metazooplankter Leptodora kindtii which controlled the abundance of herbivorous metazooplankters. We found a negative relationship between the number of bacteria and flagellates in the fishless pond, while the number of bacterivorous ciliates was suppressed by predaceous ciliates. Therefore the bacteria-grazing activity was higher in the absence of planktivorous fish.     

Effects of temporal patterning of predation threat on movement of a stream fish: evaluating an intermediate threat hypothesis

The addition of nocturnal, Hoplias malabaricus, and diurnal, Crenicichla alta, predatory fishes downstream of barrier waterfalls increases predation threat for a killifish, Rivulus hartii, in Trinidadian streams. We hypothesized that the diel patterning of predation risk would affect prey movement rates, and tested this hypothesis by comparing movement in river sites/zones containing both the nocturnal and diurnal predator with movement in river sites/zones containing only the nocturnal taxon. We evaluated this prediction in the framework of an intermediate threat hypothesis (ITH) that holds that movement will be highest at some intermediate level of threat. We marked prey fish in study sites in two watersheds of a river, each with waterfalls that divided the river into three zones: a predator absent zone (P0), a zone with one nocturnal predator (P1), and a zone with one nocturnal and one diurnal predator (P2), and tested the ITH prediction that movement will be ordered as P0<P1>P2. The single predator promoted longitudinal movement by Rivulus (P0<P1), while zones with the two predators retarded movement for small Rivulus (P1>P2) as predicted by the ITH. However, movement by larger, less vulnerable Rivulus remained elevated (P1=P2 or P2>P1). A displacement experiment in each zone found that threat tended to reduce the probability of a displaced fish reaching home, but the two predator zones did not differ from one another in their effect on this probability. Hence, the prediction that predator activity over the full 24 h diel cycle would retard movement, P2<P1, was not supported with respect to homing. Because habitat and predator communities change predictably from headwater streams to larger rivers in many lotic ecosystems, we present a conceptual model for predicting fish movement behavior along this continuum. The model posits an important role for predation threat, and the size and spacing of refuge patches, suggesting that human alterations of these factors will affect the natural movement of fish in streams.     

Direct and indirect effects of predation on mosquitofish behavior and survival

Predation can have strong direct and indirect effects on the behavior of prey. We investigated whether predation by chain pickerel (Esox niger) caused adult eastern mosquitofish (Gambusia holbrooki) to alter their habitat use and whether pickerel predation influenced survival of adult and neonate mosquitofish. The number of adult mosquitofish using the riskier of three habitats was lowest when two predators occupied the risky habitat, intermediate in the treatment with one predator, and highest when no predators occurred there. More mosquitofish neonates survived high predation treatments than treatments lacking pickerel. We conclude that pickerel predation causes adult mosquitofish to shift to refuge habitats. The pattern of neonate survival suggests that adult habitat use may create a refuge from cannibalism for neonate mosquitofish, resulting in higher neonate survival in treatments with more pickerel. Hence, pickerel predation has a direct effect on adult mosquitofish behavior and a strong indirect effect on neonate survival. Both interspecific and intraspecific predation can effect prey populations and can interact to produce important indirect effects.     

Effects of intraguild predation on resource competition.

In this work, a simple Lotka-Volterra model of intraguild predation with three species is analysed, searching for the effect of the top predator on the coexistence with its prey-competitor species. Apart from the well-known result that the intraguild prey must be superior in the competition for the shared prey in order to make coexistence possible, the magnitude of intraguild predation and the form by which the intraguild predator makes use of the intraguild prey have important consequences upon the dynamics, extending or restricting the possibilities of coexistence. These results are easily obtained by nullcline analysis. Also, some interesting results are obtained for the same model but including saturating functional response.     

Predator release of the gastropod <Emphasis Type="Italic">Cyphoma gibbosum</Emphasis> increases predation on gorgonian corals

When large, predatory fishes and invertebrates were excluded from areas of a coral reef in the Florida Keys, USA, densities of the normally rare gastropod Cyphoma gibbosum, a principal predator of gorgonian corals, increased 19-fold. Gorgonians in predator exclosures were grazed more frequently and extensively by C. gibbosum than were gorgonians in uncaged areas. In exclosures, 14% of all gorgonians showed recent predation by C. gibbosum, with 62% of the entire colony surface being removed from these attacked individuals. In areas where predators of C. gibbosum were not excluded, only 5% of gorgonians exhibited recent damage, with only 26% of the colony surface being removed from these few damaged individuals. Thus, the increases in both frequency and extent of attack combined to produce an 8× increase in gorgonian damage following removal of large predators. These patterns suggest that predators typically suppress C. gibbosum populations, that overfishing of these predators could release C. gibbosum from top-down control, and that this release will allow increased damage to gorgonian corals.     

Predation, competition, and nutrient recycling: a stoichiometric approach with multiple nutrients

A model for two competing prey species and one predator is formulated in which three essential nutrients can limit growth of all populations. Prey take up dissolved nutrients and predators ingest prey, assimilating a portion of ingested nutrients and recycling or respiring the balance. For all species, the nutrient contents of individuals vary and growth is coupled to increasing content of the limiting nutrient. This model was parameterized to describe a flagellate preying on two bacterial species, with carbon (C), nitrogen (N), and phosphorus (P) as nutrients. Parameters were chosen so that the two prey species would stably coexist without predators under some nutrient supply conditions. Using numerical simulations, the long-term outcomes of competition and predation were explored for a gradient of N:P supply ratios, varying C supply, and varying preference of the predator for the two prey. Coexistence and competitive exclusion both occurred under some conditions of nutrient supply and predator preference. As in simpler models of competition and predation these outcomes were largely governed by apparent competition mediated by the predator, and resource competition for nutrients whose effective supply was partly governed by nutrient recycling also mediated by the predator. For relatively small regions of parameter space, more complex outcomes with multiple attractors or three-species limit cycles occurred. The multiple constraints posed by multiple nutrients held the amplitudes of these cycles in check, limiting the influence of complex dynamics on competitive outcomes for the parameter ranges explored.