Cascading effects of predator diversity and omnivory in a marine food web

Over‐harvesting, habitat loss and exotic invasions have altered predator diversity and composition in a variety of communities which is predicted to affect other trophic levels and ecosystem functioning. We tested this hypothesis by manipulating predator identity and diversity in outdoor mesocosms that contained five species of macroalgae and a macroinvertebrate herbivore assemblage dominated by amphipods and isopods. We used five common predators including four carnivores (crabs, shrimp, blennies and killifish) and one omnivore (pinfish). Three carnivorous predators each induced a strong trophic cascade by reducing herbivore abundance and increasing algal biomass and diversity. Surprisingly, increasing predator diversity reversed these effects on macroalgae and altered algal composition, largely due to the inclusion and performance of omnivorous fish in diverse predator assemblages. Changes in predator diversity can cascade to lower trophic levels; the exact effects, however, will be difficult to predict due to the many complex interactions that occur in diverse food webs.     

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Cascading effects of predatory fish on the composition of benthic algae in high‐altitude streams

Cascading effects of predators can affect ecosystem properties by changing plant biomass, distribution and assemblage composition. Using data from field surveys and whole‐stream experiments we tested the hypothesis that predatory trout change assemblage composition of benthic algae in high‐elevation streams mediated by grazer behavior. Field surveys revealed that the taxonomic composition of algal assemblages differed significantly between streams that contained trout and those that were fishless; but comparisons of palatable versus unpalatable algal taxa between fish and fishless streams were equivocal because of high natural variability. Therefore, we tested for a behavioral (non‐consumptive) trophic cascade experimentally by adding brook trout chemical cues to six naturally fishless streams for 25 days and compared responses of grazers and algae to six reference streams without fish cues added. Algal response variables included rates of change in the abundance of three physiognomic categories, from most palatable (attached erect and prostrate diatoms) to least palatable (non‐diatoms), as determined from food selectivity analyses of the most common grazers (mayflies and caddisflies). Fish cues did not affect the mean densities or changes in densities of total grazers or any individual grazer species. However, in streams where fish cues were added, rates of accrual of attached erect diatoms, which was the preferred algal type for the grazer most vulnerable to trout predation (Baetis), were higher and their densities increased significantly faster with increasing densities of this grazer species than in reference streams. Results of his experiment support the hypothesis that predator induced suppression of grazer foraging behavior, rather than cascading effects of top predators on grazer density, may contribute to variation in the composition of algal assemblages among streams by allowing proliferation of most palatable algal species.     

Environmental context determines multi‐trophic effects of consumer species loss

Loss of biodiversity and nutrient enrichment are two of the main human impacts on ecosystems globally, yet we understand very little about the interactive effects of multiple stressors on natural communities and how this relates to biodiversity and ecosystem functioning. Advancing our understanding requires the following: (1) incorporation of processes occurring within and among trophic levels in natural ecosystems and (2) tests of context‐dependency of species loss effects. We examined the effects of loss of a key predator and two groups of its prey on algal assemblages at both ambient and enriched nutrient conditions in a marine benthic system and tested for interactions between the loss of functional diversity and nutrient enrichment on ecosystem functioning. We found that enrichment interacted with food web structure to alter the effects of species loss in natural communities. At ambient conditions, the loss of primary consumers led to an increase in biomass of algae, whereas predator loss caused a reduction in algal biomass (i.e. a trophic cascade). However, contrary to expectations, we found that nutrient enrichment negated the cascading effect of predators on algae. Moreover, algal assemblage structure varied in distinct ways in response to mussel loss, grazer loss, predator loss and with nutrient enrichment, with compensatory shifts in algal abundance driven by variation in responses of different algal species to different environmental conditions and the presence of different consumers. We identified and characterized several context‐dependent mechanisms driving direct and indirect effects of consumers. Our findings highlight the need to consider environmental context when examining potential species redundancies in particular with regard to changing environmental conditions. Furthermore, non‐trophic interactions based on empirical evidence must be incorporated into food web‐based ecological models to improve understanding of community responses to global change.     

Food chain length and omnivory determine the stability of a marine subtidal food web

1. Using a subtidal marine food web as a model system, we examined how food chain length (predators present or absent) and the prevalence of omnivory influenced temporal stability (and its components) of herbivores and plants. We held the density of top predators constant but manipulated their identity to generate a gradient in omnivory prevalence. 2. We measured temporal stability as the inverse of the coefficient of variation of abundance over time. Predators and omnivory could influence temporal stability through effects on abundance (the 'abundance' effect), summed variance across taxa (the 'portfolio effect') or summed covariances among taxa (the 'covariance effect'). 3. We found that increasing food chain length by predator addition destabilized aggregate herbivore abundance through their cascading effects on abundances. Thus, predators destabilized herbivores through the overyielding effect. We also found that the stability of herbivore abundance and microalgae declined with increasing prevalence of omnivory among top predators. Aggregate macroalgae was not affected, but the stability of one algal taxon increased with the prevalence of omnivory. 4. Our results suggest that herbivores are more sensitive than plants to changes in food web structure because of predator additions by invasion or deletions such as might occur via harvesting and habitat loss.     

Trophic cascades in a temperate seagrass community

We assessed the relative importance of bottom–up and top–down processes in structuring an eelgrass community in Sweden, a system impacted both by eutrophication and overfishing. Using artificial seagrass as substrate, we manipulated nutrient levels and predator abundance in a full‐factorial cage‐experiment. The results revealed a seagrass community dominated by strong top–down processes controlling the aggregate biomass of mesograzers and macroalgae. In the absence of predators the large amphipod Gammarus locusta became very abundant resulting in a leaf community with low biomass of algae and smaller mobile fauna. One enclosed gobid fish predator reduced the abundance of adult G. locusta by >90%, causing a three to six times increase in the biomass of algae, smaller mesograzers and meiofauna. Numerous small predators in uncaged habitats reduced the biomass of G. locusta and other mesograzers by >95% in comparison to the fish treatment, further increasing the biomass of epiphytic algae and meiofauna. Although water column nutrient enrichment caused a temporal bloom of the filamentous macroalgae Ulva spp., no significant nutrient‐effects were found on the algal community at the end of the experiment. The only lasting nutrient‐effect was a significant increase in the biomass of G. locusta, but only in the absence of ambient predators. These results demonstrate that mesograzers can respond to enhanced food supply, increase their biomass and control the algal growth when predation rates are low. However, in the assessed system, high predation rates appear to make mesograzers functionally extinct, causing a community‐wide trophic cascade that promotes the growth of ephemeral algae. This top–down effect could penetrate down, despite a complex food‐web because the interaction strength in the community was strongly skewed towards two functionally dominant algal and grazer species that were vulnerable to consumption. These results indicate that overexploitation of gadoid fish may be linked to increased macroalgal blooms and loss of eelgrass in the area through a trophic cascade affecting the abundance of mesograzers.     

Herbivore species richness, composition and community structure mediate predator richness effects and top-down control of herbivore biomass

Changes in predator species richness can have important consequences for ecosystem functioning at multiple trophic levels, but these effects are variable and depend on the ecological context in addition to the properties of predators themselves. Here, we report an experimental study to test how species identity, community attributes, and community structure at the herbivore level moderate the effects of predator richness on ecosystem functioning. Using mesocosms containing predatory insects and aphid prey, we independently manipulated species richness at both predator and herbivore trophic levels. Community structure was also manipulated by changing the distribution of herbivore species across two plant species. Predator species richness and herbivore species richness were found to negatively interact to influence predator biomass accumulation, an effect which is hypothesised to be due to the breakdown of functional complementarity among predators in species-rich herbivore assemblages. The strength of predator suppression of herbivore biomass decreased as herbivore species richness and distribution across host plants increased, and positive predator richness effects on herbivore biomass suppression were only observed in herbivore assemblages of relatively low productivity. In summary, the study shows that the species richness, productivity and host plant distribution of prey communities can all moderate the general influence of predators and the emergence of predator species richness effects on ecosystem functioning.     

Cascading effects of predatory fish exclusion on the detritus-based food web of a lake littoral zone (Lake Vico, central Italy)

An exclosure experiment was carried out in the reed-dominated littoral zone of a volcanic lake (Lake Vico, central Italy) to test whether the impact of predatory fish on benthic invertebrates cascades on fungal colonisation and breakdown of leaf detritus. The abundance, biomass, and Shannon diversity index of the invertebrate assemblage colonising Phragmites australis leaf packs placed inside: (1) full-exclosure cages, (2) cages allowing access only to small-sized fish predators, and (3) cageless controls, were monitored over a 45-day period together with the mass loss and associated fungal biomass of leaf packs. The species composition of the fungal assemblage was further assessed at the end of the manipulation. In general, invertebrate predators did not show any significant response to fish exclusion, either on a trophic guild or on a single taxon level. In contrast, the exclusion of large predatory fish induced a diverse spectrum of changes in the abundance and population size-structure of dominant detritivore taxa, ultimately increasing the biomass and Shannon diversity index of the whole detritivorous guild. These changes corresponded with significant variations in leaf detritus decay rates as well as in the biomass and assemblage structure of associated fungal colonisers. Our experimental findings provide evidence that in Lake Vico effects of fish predators on invertebrate detritivores influence the fungal conditioning and breakdown of the detrital substrate. We conclude that in lacustrine littoral zones predator-driven constraints may structure lower trophic levels of detritus-based food webs and affect the decomposition of leaf detritus originated from the riparian vegetation.     

Are impacts of an exotic predator on a stream food web influenced by disturbance history?

Predatory species have been introduced to habitats spanning a wide range of environmental conditions. To better understand the consequences of predation in natural communities we need to examine how variations in abiotic factors modify the influence of predation. The effects of introduced predators may vary amongst habitats if natural disturbance affects the abundance and taxonomic composition of consumers and their resources, or the predator alters recolonisation after disturbance. We tested whether a bed-moving disturbance altered subsequent interactions involving native and introduced predatory fish, invertebrate grazers and algae in experimental channels within a New Zealand stream. Disturbance reduced the abundance of invertebrates by 84%, and induced mortality of Conoesucidae caddisflies. However, the relative abundance of taxa changed little immediately following the disturbance. Invertebrate communities recovered following disturbance in fishless channels and those with native galaxiids (Galaxias vulgaris), and were almost indistinguishable from undisturbed fishless controls after 2 weeks. Invertebrate abundance declined and algal abundance increased in channels with exotic brown trout (Salmo trutta) and their effect was strongest in previously disturbed channels. However, predators and disturbance only had interactive effects on grazer emigration rates. Trout affected grazers through direct consumption (e.g. Conoesucidae caddisflies), and induced higher emigration rates of grazers from channels via drift (e.g. the mayfly Deleatidium). The effects of predatory trout and galaxiids combined differed in disturbed and stable channels. The observed combined effects of predatory trout and galaxiids on invertebrate grazers were lower than expected in stable channels partly due to low emigration rates of Conoesucidae, whereas emigration of grazers was higher than expected in the disturbed channels. The biomass of algae was higher than expected in disturbed channels with both predators. Collectively, our results indicate that predator substitutability and the non-lethal effects of introduced predators varied depending on disturbance history, but their effects on the biomass of grazers and algae did not.     

Ecological factors associated with the strength of trophic cascades in streams

Trophic cascades are extensively documented in nature, but they are also known to vary widely in strength and frequency across ecosystems. Therefore, much effort has gone into understanding which ecological factors generate variation in cascade strength. To identify which factors covary with the strength of cascades in streams, we performed a concurrent experiment across 17 streams throughout the Sierra Nevada Mountains. We eliminated top consumers from experimental substrates using electrical exclusions and compared the strength of indirect effects of consumers on the biomass of primary producers relative to control patches. In each stream we 1) classified the dominant invertebrate herbivores according to life‐history traits that influence their susceptibility to predators, 2) determined the abundance and diversity of algae and herbivores, and 3) measured production‐to‐biomass ratios (P:B) of the stream biofilm. This allowed us to assess three common predictions about factors thought to influence the strength of trophic cascades: cascade strength 1) is weaker in systems dominated by herbivores with greater ability to evade or defend against predators, 2) is stronger in systems characterized by low species diversity, and 3) increases with increasing producer P:B. When averaged across all streams, the indirect effect of predators increased the biomass of periphyton by a mean 60%. However, impacts of predators on algae varied widely, ranging from effects that exacerbated algal loss to herbivores, to strong cascades that increased algal biomass by 4.35 times. Cascade strength was not related to herbivore traits or species diversity, but decreased significantly with increasing algal diversity and biofilm P:B in a stream. Partial regression analyses suggested that the relationship between cascade strength and algal diversity was spurious, and that the only significant covariate after statistically controlling for cross‐correlations was algal P:B. Our study contributes to the ongoing debate about why trophic cascade strength varies in nature and is useful because it eliminates factors that have no potential to explain variation in cascades within these stream ecosystems.     

Top–down limits on prey populations may be more severe in larger prey species,despite having fewer predators

Variation in the vulnerability of herbivore prey to predation is linked to body size, yet whether this relationship is size‐nested or size‐partitioned remains debated. If size‐partitioned, predators would be focused on prey within their preferred prey size range. If size‐nested, smaller prey species should become increasingly more vulnerable because increasingly more predators are capable of catching them. Yet, whether either of these strategies manifests in top–down prey population limitation would depend both on the number of potential predator species as well as the total mortality imposed. Here we use a rare ecosystem scale ‘natural experiment’ comparing prey population dynamics between a period of intense predator persecution and hence low predator densities and a period of active predator protection and population recovery. We use three decades of data on herbivore abundance and distribution to test the role of predation as a mechanism of population limitation among prey species that vary widely in body size. Notably, we test this within one of the few remaining systems where a near‐full suite of megaherbivores occur in high density and are thus able to include a thirtyfold range in herbivore body size gradient. We test whether top–down limitation on prey species of particular body size leads to compositional shifts in the mammalian herbivore community. Our results support both size‐nested and size‐partitioning predation but suggest that the relative top–down limiting impact on prey populations may be more severe for intermediate sized species, despite having fewer predators than small species. In addition we show that the gradual recovery of predator populations shifted the herbivore community assemblage towards large‐bodied species and has led to a community that is strongly dominated by large herbivore biomass.     

Herbivore and predator diversity interactively affect ecosystem properties in an experimental marine community

Interacting changes in predator and prey diversity likely influence ecosystem properties but have rarely been experimentally tested. We manipulated the species richness of herbivores and predators in an experimental benthic marine community and measured their effects on predator, herbivore and primary producer performance. Predator composition and richness strongly affected several community and population responses, mostly via sampling effects. However, some predators survived better in polycultures than in monocultures, suggesting complementarity due to stronger intra- than interspecific interactions. Predator effects also differed between additive and substitutive designs, emphasizing that the relationship between diversity and abundance in an assemblage can strongly influence whether and how diversity effects are realized. Changing herbivore richness and predator richness interacted to influence both total herbivore abundance and predatory crab growth, but these interactive diversity effects were weak. Overall, the presence and richness of predators dominated biotic effects on community and ecosystem properties.     

Top predator control of plant biodiversity and productivity in an old-field ecosystem

Abstract Predators can have strong indirect effects on plants by altering the way herbivores impact plants. Yet, many current evaluations of plant species diversity and ecosystem function ignore the effects of predators and focus directly on the plant trophic level. This report presents results of a 3‐year field experiment in a temperate old‐field ecosystem that excluded either predators, or predators and herbivores and evaluated the consequence of those manipulations on plant species diversity (richness and evenness) and plant productivity. Sustained predator and predator and herbivore exclusion resulted in lower plant species evenness and higher plant biomass production than control field plots representing the intact natural ecosystem. Predators had this diversity‐enhancing effect on plants by causing herbivores to suppress the abundance of a competitively dominant plant species that offered herbivores a refuge from predation risk.     

Keystone predator effects and grazer control of planktonic primary production

If prey species exhibit trade-offs in their ability to utilize resources versus their ability to avoid predation, predators can facilitate prey turnover along gradients of productivity, shifting dominance from edible to inedible prey (the keystone predator effect). I tested this model under controlled, laboratory conditions, using a model aquatic system composed of zooplankton as the top consumer, a diverse community of algae as prey, and nutrients as basal resources. Nutrient manipulations (low and high) were crossed with presence–absence of zooplankton. Results supported theoretical predictions. Algal biomass increased in response to enrichment regardless of predator presence/absence. However, predators and nutrients had an interactive effect on algal biomass and size structure. At the low nutrient level, algal-prey were dominated by edible forms and attained similar biomass regardless of zooplankton presence/absence. At the high level of enrichment, presence of zooplankton favored higher levels of algal biomass and shifted dominance to large, inedible taxa. At the termination of the experiment, I performed a series of lab-based assays on the resultant algal community in order to quantify trade-offs among algal size classes in maximal population growth rates (as a measure of competitive ability for nutrients) and susceptibility to zooplankton grazing. Assays provided support for a size-based keystone trade-off. Small size classes of algae displayed higher maximal growth rates but were more susceptible to grazing effects. Large size classes were protected from grazing but showed low rates of population growth in response to enrichment.     

Fish predation regimes modify benthic diatom community structures: Experimental evidence from an in situ mesocosm study

Diatoms are important primary producers in shallow water environments. Few studies have assessed the importance of biological interactions in structuring these communities. In the present study, benthic diatom community structure in relation to manipulated food webs was assessed using in situ mesocosms, whereby predator‐free environments and environments comprising two different fish species were assessed. Zooplankton abundance, settled algal biomass and the diatom community were monitored over a 12‐day period across each of the three trophic scenarios. Differences among treatments over time were observed in zooplankton abundances, particularly copepods. Similarly, the benthic diatom community structure changed significantly over time across the three trophic treatments. However, no differences in total algal biomass were found among treatments. This was likely the result of non‐diatom phytoplankton contributions. We propose that the benthic diatom community structure within the mesocosms was influenced by trophic cascades and potentially through direct consumption by the fish. The study highlights that not only are organisms at the base of the food web affected by predators at the top of the food web, but that predator identity is potentially an important consideration for predator–prey interaction outcomes with consequences for multiple trophic levels.     

Spatial refuge from intraguild predation: implications for prey suppression and trophic cascades

The ability of predators to elicit a trophic cascade with positive impacts on primary productivity may depend on the complexity of the habitat where the players interact. In structurally-simple habitats, trophic interactions among predators, such as intraguild predation, can diminish the cascading effects of a predator community on herbivore suppression and plant biomass. However, complex habitats may provide a spatial refuge for predators from intraguild predation, enhance the collective ability of multiple predator species to limit herbivore populations, and thus increase the overall strength of a trophic cascade on plant productivity. Using the community of terrestrial arthropods inhabiting Atlantic coastal salt marshes, this study examined the impact of predation by an assemblage of predators containing Pardosa wolf spiders, Grammonota web-building spiders, and Tytthus mirid bugs on herbivore populations (Prokelisia planthoppers) and on the biomass of Spartina cordgrass in simple (thatch-free) and complex (thatch-rich) vegetation. We found that complex-structured habitats enhanced planthopper suppression by the predator assemblage because habitats with thatch provided a refuge for predators from intraguild predation including cannibalism. The ultimate result of reduced antagonistic interactions among predator species and increased prey suppression was enhanced conductance of predator effects through the food web to positively impact primary producers. Behavioral observations in the laboratory confirmed that intraguild predation occurred in the simple, thatch-free habitat, and that the encounter and capture rates of intraguild prey by intraguild predators was diminished in the presence of thatch. On the other hand, there was no effect of thatch on the encounter and capture rates of herbivores by predators. The differential impact of thatch on the susceptibility of intraguild and herbivorous prey resulted in enhanced top-down effects in the thatch-rich habitat. Therefore, changes in habitat complexity can enhance trophic cascades by predator communities and positively impact productivity by moderating negative interactions among predators.     

A comparison of algal,macroinvertebrate, and fish assemblage indices for assessing low-level nutrient enrichment in wadeable Ozark streams

Biotic indices for algae, macroinvertebrates, and fish assemblages can be effective for monitoring stream enrichment, but little is known regarding the value of the three assemblages for detecting perturbance as a consequence of low-level nutrient enrichment. In the summer of 2006, we collected nutrient and biotic samples from 30 wadeable Ozark streams that spanned a nutrient-concentration gradient from reference to moderately enriched conditions. Seventy-three algal metrics, 62 macroinvertebrate metrics, and 60 fish metrics were evaluated for each of the three biotic indices. After a group of candidate metrics had been identified with multivariate analysis, correlation procedures and scatter plots were used to identify the four metrics having strongest relations to a nutrient index calculated from log transformed and normalized total nitrogen and total phosphorus concentrations. The four metrics selected for each of the three biotic indices were: algae—the relative abundance of most tolerant diatoms, the combined relative abundance of three species of Cymbella, mesosaprobic algae percent taxa richness, and the relative abundance of diatoms that are obligate nitrogen heterotrophs; macroinvertebrate—the relative abundance of intolerant organisms, Baetidae relative abundance, moderately tolerant taxa richness, and insect biomass; fish—herbivore and detritivore taxa richness, pool species relative abundance, fish catch per unit effort, and black bass (Micropterus spp.) relative abundance.All three biotic indices were negatively correlated to nutrient concentrations but the algal index had a higher correlation (rho = ?0.89) than did the macroinvertebrate and fish indices (rho = ?0.63 and ?0.58, respectively). Biotic index scores were lowest and nutrient concentrations were highest for streams with basins having the highest poultry and cattle production. Because of the availability of litter for fertilizer and associated increases in grass and hay production, cattle feeding capacity increases with poultry production. Studies are needed that address the synergistic effect of poultry and cattle production on Ozark streams in high production areas before ecological risks can be adequately addressed.     

Effects of herbivore exclusion and nutrient enrichment on coral reef macroalgae and cyanobacteria

Although phase shifts on coral reefs from coral-dominated to algal-dominated communities have been attributed to the effects of increased nutrient availability due to eutrophication and reduced herbivore abundance due to overfishing and disease, these factors have rarely been manipulated simultaneously. In addition, few studies have considered the effects of these factors on benthic, filamentous cyanobacteria (blue-green algae) as well as macroalgae. We used a combination of herbivore-exclusion cages and nutrient enrichment to manipulate herbivore abundance and nutrient availability, and measured the impacts of these treatments on macroalgal and cyanobacterial community structure. In the absence of cages, surface cover of the cyanobacterium Tolypothrix sp. decreased, while surface cover of the cyanobacteria Oscillatoria spp. increased. Cyanobacterial cover decreased in partial cages, and Tolypothrix sp. cover decreased further in full cages. Lower cyanobacterial cover and biomass were correlated with higher macroalgal cover and biomass. Dictyota bartayresiana dominated the partial cages, while Padina tenuis and Tolypiocladia glomerulata recruited into the full cages. Palatability assays demonstrated that herbivore-exclusion shifted macroalgal species composition from relatively unpalatable to relatively palatable species. Nutrient enrichment interacted with herbivore exclusion to increase the change in cover of D. bartayresiana in the uncaged and fully caged plots, but did not affect the final biomass of D. bartayresiana among treatments. Nutrient enrichment did not significantly affect the cover or biomass of any other taxa. These results stress the critical role of herbivory in determining coral reef community structure and suggest that the relative palatabilities of dominant algae, as well as algal growth responses to nutrient enrichment, will determine the potential for phase shifts to algal-dominated communities.     

The effect of two scales of habitat architecture on benthic grazing in a river

1. This experiment studied the effects of differing levels of the complexity of substratum architecture at two spatial scales on the distribution and abundance of benthic algae and invertebrates, and the strength of the trophic interaction between invertebrate grazers and algae. Some estimates of the effects on invertebrate colonization rates were also made. 2. Four levels of microhabitat architectural complexity were created using artificial substrata (clay tiles) and placed in Mountain River, Tasmania, in two riffle types (bedrock and boulder-cobble) of differing large-scale substratum complexity. After a colonization period, invertebrate grazers were removed from half the tiles to measure the effects of grazing. Invertebrates on the tiles were also counted and identified. At the end of the experiment, algae were removed from the tiles and analysed for chlorophyll a. 3. Invertebrate grazers did not reduce algal biomass during the experiment, and microhabitat-scale architecture influenced algal biomass more strongly than riffle-scale architecture. Highly complex microhabitat architecture increased algal biomass by providing more surface area, but once standardized for surface area, algal biomass decreased as the complexity of microhabitat architecture increased. 4. Microhabitat-scale architecture was also predominant in determining invertebrate density and the identity of the dominant grazer species. In contrast to algal biomass, invertebrate densities and species density increased with the complexity of microhabitat architecture, suggesting that refuges from flow (and possibly predation) were as important to river invertebrates as the distribution of their food source. 5. Riffle-scale architecture had some effect on the colonization of two slow-moving grazer taxa, but, overall, the colonization processes of slow-moving grazers were determined mostly by the complexity of microhabitat-scale architecture.     

Detriments to post-bleaching recovery of corals

Predicting the response of coral reefs to large-scale mortality induced by climate change will depend greatly on the factors that influence recovery after bleaching events. We experimentally transplanted hard corals from a shallow reef with highly variable seawater temperature (23–36°C) to three unfished marine parks and three fished reefs with variable coral predator abundance and benthic cover. The transplanted corals were fragmented colonies collected from a reef that was relatively undisturbed by the 1997–1998 warm-water temperature anomaly, one of the most extreme thermal events of the past century, and it was assumed that they would represent corals likely to succeed in the future temperature environment. We examined the effects of four taxa, two fragment sizes, an acclimation period, benthic cover components, predators and tourists on the survival of the coral fragments. We found the lowest survival of transplants occurred in the unfished marine parks and this could be attributed to predation and not tourist damage. The density of small coral recruits approximately 6 months after the spawning season was generally moderate (~40–60/m²), and not different on fished and unfished reefs. Coral recovery between 1998 and 2002 was variable (0–25%), low (mean of 6.5%), and not different between fished and unfished reefs. There was high variability in coral mortality among the three unfished areas despite low variation in estimates of predator biomass, with the highest predation occurring in the Malindi MNP, a site with high coralline algal cover. Stepwise multiple regression analysis with 14 variables of coral predators and substratum showed that coralline algae was positively, and turf algae negatively associated with mortality of the transplants, with all other variables being statistically insignificant. This suggests that alternate food resources and predator choices are more important than predator biomass in determining coral survival. Nonetheless, large predatory fish in areas dominated by coralline algae may considerably retard recovery of eurythermal corals. This will not necessarily retard total hard coral recovery, as other more predator-tolerant taxa can recover. Based on the results, global climate change will not necessarily favor eurythermal over stenothermal coral taxa in remote or unfished reefs, where predation is a major cause of coral mortality.