Strength Of Top-down Control Research Articles

Top-down versus bottom-up: Grazing and upwelling regime alter patterns of primary productivity in a warm-temperate system.

Community structure is driven by the interaction of physical processes and biological interactions that can vary across environmental gradients and the strength of top-down control is expected to vary along gradients of primary productivity. In coastal marine systems, upwelling drives regional resource availability through the bottom-up effect of nutrient subsidies. This alters rates of primary production and is expected to alter algae-herbivore interactions in rocky intertidal habitats. Despite the potential for upwelling to alter these interactions, the interaction of upwelling and grazing pressure is poorly understood, particularly for warm-temperate systems. Using in situ herbivore exclusion experiments replicated across multiple upwelling regimes, we investigated the effects of both grazing pressure, upwelling, and their interactions on the sessile invertebrate community and biomass of macroalgal communities in a warm-temperate system. The sessile invertebrate cover showed indirect effects of grazing, being consistently low where algal biomass was high at upwelling sites and at nonupwelling sites when grazers were excluded. The macroalgal cover was greater at upwelling sites when grazers were excluded and there was a strong effect of succession throughout the experimental period. Grazing effects were greater at upwelling sites, particularly during winter months. There was a nonsignificant trend toward greater grazing pressure on early than later successional stages. Our results show that the positive bottom-up effects of nutrient supply on algal production do not overwhelm top-down control in this warm-temperate system but do have knock-on consequences for invertebrates that compete with macroalgae for space. We speculate that global increases in air and sea surface temperatures in warm-temperate systems will promote top-down effects in upwelling regions by increasing herbivore metabolic and growth rates.

What Drives the Assembly of Plant-associated Protist Microbiomes? Investigating the Effects of Crop Species, Soil Type and Bacterial Microbiomes

In a field experiment we investigated the influence of the environmental filters soil type (i.e. three contrasting soils) and plant species (i.e. lettuce and potato) identity on rhizosphere community assembly of Cercozoa, a dominant group of mostly bacterivorous soil protists. Plant species (14%) and rhizosphere origin (vs bulk soil) with 13%, together explained four times more variation in cercozoan beta diversity than the three soil types (7% explained variation). Our results clearly confirm the existence of plant species-specific protist communities. Network analyses of bacteria-Cercozoa rhizosphere communities identified scale-free small world topologies, indicating mechanisms of self-organization. While the assembly of rhizosphere bacterial communities is bottom-up controlled through the resource supply from root (secondary) metabolites, our results support the hypothesis that the net effect may depend on the strength of top-down control by protist grazers. Since grazing of protists has a strong impact on the composition and functioning of bacteria communities, protists expand the repertoire of plant genes by functional traits, and should be considered as 'protist microbiomes' in analogy to 'bacterial microbiomes'.

Can young-of-the-year invasive fish keep up with young-of-the-year native fish? A comparison of feeding rates between invasive sticklebacks and whitefish.

Invasion of non‐native species might alter food web structure and the strength of top‐down control within lake ecosystems. As top‐down control exerted by fish populations is often dominated by young of the year fish, the impact of new fish species might depend on the feeding rates of the juvenile fish. Here we provide comparative analyses of feeding rates of juvenile whitefish (Coregonus wartmanni) – a native and specialised planktivore and an invasive generalist (sticklebacks, Gasterosteus aculeatus). We studied feedings rates of whitefish and sticklebacks in aquaria experiments using 2 cm to 8 cm fish feeding on seven zooplankton species common to Lake Constance. As whitefish hatch several months earlier than sticklebacks, 0+ whitefish are larger than 0+ sticklebacks throughout the year and hence are predicted to have higher feeding rates on especially large zooplankton species. We show that sticklebacks as small as 2 cm were able to feed on the largest zooplankton species of Lake Constance. Further, stickleback feeding rates were similar to both the same size 0+ whitefish and the larger 0+ whitefish co‐occurring with smaller 0+ sticklebacks. Hence, 0+ sticklebacks will compete with 0+ whitefish for the same zooplankton species, therefore the invasion of sticklebacks is unlikely to change the relative feeding pressure by individual 0+ fish on zooplankton species.

Multi-scale mosaics in top-down pest control by ants from natural coffee forests to plantations.

While top-down control plays an important role in shaping both natural and agricultural food webs, we lack insights into how top-down control effects vary across spatial scales. We used a multi-scale survey of top-down control of coffee pests and diseases by arboreal ants to examine if colony location creates a small-scale mosaic in top-down control around trees and if the strength of that control varies between sites at the landscape scale. We investigated pest and disease levels on coffee shrubs at different distances from shade trees with and without a Crematogaster spp. ant colony in 59 sites along a coffee management intensity gradient in southwestern Ethiopia. Within sites, ants significantly suppressed herbivory and coffee leaf rust at distances less than 10m from nesting trees. Top-down control varied between sites, with stronger top-down control of free-feeding herbivory near ant colonies at sites with lower management intensity and stronger top-down control of a skeletonizer at sites with higher canopy cover. We conclude that the strength of top-down control by ants is highly heterogeneous across spatial scales, as a consequence of the biology of the predator at the small scale and herbivore density or changes in herbivore-ant interactions at the landscape scale.

Biogeographic context dependence of trophic cascade strength in bromeliad food webs.

Ecosystem functions and the biomass of lower trophic levels are frequently controlled by predators. The strength of top-down control in these trophic cascades can be affected by the identity and diversity of predators, prey, and resources, as well as environmental conditions such as temperature, moisture, and nutrient loading, which can all impact interaction strength between trophic levels. Few studies have been able to replicate a complete community over a large geographic area to compare the full trophic cascade in a manipulative experiment. Here, we identify geographic dependency in trophic cascade strength, and the driving factors and specific mechanisms behind it, by combining geographically replicated experiments with a novel approach of community analogues of common garden and transplant experiments. We studied a predator-detritivore-detritus food web in bromeliads in Puerto Rico, Costa Rica, and Brazil. We found that interaction strengths between resources, consumers, and predators were strongly site-specific, but the exact mechanism differed between trophic levels. Large bodied predators created strong interaction strengths between predator and consumer trophic levels, reducing consumer abundance regardless of the geographic location, whereas small-bodied predators created weak interactions with no impact on consumer abundances in any site. In contrast, the interaction strength between consumers and resources varied among sites, depending on the dominant species of leaf detritus. More labile leaf species in Costa Rica created a strong consumer-resource interaction and therefore strong trophic cascade, whereas tougher leaf species in Brazil created a weak consumer-resource interaction, and an overall weaker trophic cascade. Our study highlights the importance of replicating experiments over geographic scales to understand general patterns of ecological processes.

Landscape patterns in top-down control of decomposition: omnivory disrupts a tropical detrital-based trophic cascade.

Detrital-based trophic cascades are often considered weak or absent in tropical stream ecosystems because of the prevalence of omnivorous macroconsumers and the dearth of leaf-shredding insects. In this study, we isolate top-down effects of three macroconsumer species on detrital processing in headwater streams draining Trinidad's northern mountains. We separated effects of different macroconsumers by experimentally manipulating their temporal access to isolated benthic habitat over the diel cycle. We found no evidence that omnivorous macroconsumers, including a freshwater crab (Eudaniela garmani) and guppy (Poecilia reticulata), increased leaf decomposition via consumption. By contrast, above a waterfall excluding guppies, the insectivorous killifish, Anablepsoides hartii, reduced the biomass of the leaf-shredding insect Phylloicus hansoni 4-fold, which consequently reduced leaf decomposition rates 1.6-fold. This detrital cascade did not occur below the barrier waterfall, where omnivorous guppies join the assemblage and reduce killifish densities; here killifish had no significant effects on Phylloicus or decomposition rates. These patterns of detrital processing were also observed in upstream-downstream comparisons in a landscape study across paired reaches of six streams. Above waterfalls, where killifish were present, but guppies absent, leaf decomposition rates and Phylloicus biomass were 2.5- and ~35-fold lower, respectively, compared to measurements below waterfalls. Moreover, the strength of top-down control by killifish is reflected by the 20- and 5-fold reductions in variability (±SE) surrounding mean Phylloicus biomass and leaf decomposition rates in upstream relative to downstream reaches where no top-down control was detected. Findings show a clear, detrital-based trophic cascade among killifish, a leaf-shredding insect, and leaf decomposition rates. Results also show how omnivorous guppies disrupt this cascade by depressing killifish densities, thereby releasing invertebrate shredders from predation, and significantly increasing decomposition rates. Moreover, this combination of direct and indirect trophic interactions drives patterns in decomposition rates in stream networks at a landscape scale, resulting in significantly lower rates of decomposition above vs. below barrier waterfalls. Our findings reveal that omnivory can result in significant indirect effects on a key ecosystem process, illustrating the importance of these hidden trophic pathways in detrital-based systems and suggesting that resource control in tropical systems may be even more complex than previously envisioned.

Accounting for activity respiration results in realistic trophic transfer efficiencies in allometric trophic network (ATN) models

Allometric trophic network (ATN) models offer high flexibility and scalability while minimizing the number of parameters and have been successfully applied to investigate complex food web dynamics and their influence on food web diversity and stability. However, the realism of ATN model energetics has never been assessed in detail, despite their critical influence on dynamic biomass and production patterns. Here, we compare the energetics of the currently established original ATN model, considering only biomass-dependent basal respiration, to an extended ATN model version, considering both basal and assimilation-dependent activity respiration. The latter is crucial in particular for unicellular and invertebrate organisms which dominate the metabolism of pelagic and soil food webs. Based on metabolic scaling laws, we show that the extended ATN version reflects the energy transfer through a chain of four trophic levels of unicellular and invertebrate organisms more realistically than the original ATN version. Depending on the strength of top-down control, the original ATN model yields trophic transfer efficiencies up to 71% at either the third or the fourth trophic level, which considerably exceeds any realistic values. In contrast, the extended ATN version yields realistic trophic transfer efficiencies ≤ 30% at all trophic levels, in accordance with both physiological considerations and empirical evidence from pelagic systems. Our results imply that accounting for activity respiration is essential for consistently implementing the metabolic theory of ecology in ATN models and for improving their quantitative predictions, which makes them more powerful tools for investigating the dynamics of complex natural communities.

Community structure and biovolume size spectra of mesozooplankton in the Pearl River estuary

The Pearl River estuary is the second largest estuary in China. This study reported the species composition and size structure of mesozooplankton in the upper and lower Pearl River estuary in the dry and wet seasons. The normalized biovolume size spectrum of mesozooplankton was constructed to analyze the trophic structure of the community. Copepods were the numerically predominant mesozooplankton group at all sampling stations. However, when converted into biovolume, medusae were dominant in the upper estuary in the dry season and Chaetognatha were dominant in the lower estuary in the wet season. There were apparent seasonal and spatial variations in the normalized biovolume size spectrum characteristics in the Pearl River estuary. In the dry season, the average fitted regression line of the normalized biovolume size spectrum had a slope of −1.02 in the upper estuary and −0.88 in the lower estuary. In the wet season, the average fitted regression line of the normalized biovolume size spectrum had a slope of −0.32 in the lower estuary, and no significant regression of the normalized biovolume size spectrum was found in the upper estuary. It is suggested that the mesozooplankton community was unstable in the upper estuary due to the strong freshwater perturbation in the wet season. The size diversity and normalized biovolume size spectrum slope indicated that the strength of top-down control and trophic efficiency was highest in the lower estuary in the wet season.

Reciprocal feedbacks between spatial subsidies and reserve networks in coral reef meta-ecosystems.

Top-down processes such as predation and herbivory have been shown to control the dynamics of communities across a range of ecosystems by generating trophic cascades. However, theory is only beginning to describe how these local trophic processes interact with spatial subsidies in the form of material (nutrient, detritus) transport and organismal dispersal to (1) shape the structure of interconnected (meta-) ecosystems and (2) determine their optimal management via reserve networks. Here, we develop a meta-ecosystem model to understand how the reciprocal feedbacks between spatial subsidies and reserve networks modulate the importance of top-down control in a simple herbivorous fish-macroalgae-coral system. We show that in large and isolated reserve networks where connectivity between protected and unprotected areas is limited, spatial subsidies remain largely confined to reserves. This retention of spatial subsidies promotes the top-down control of corals and macroalgae by herbivores inside reserves but reduces it outside reserves. Conversely, in small and aggregated reserves where connectivity between protected and unprotected areas is high, the spillover of spatial subsidies causes a reduction in top-down control of corals and macroalgae by herbivores inside reserves and an increase in the strength of top-down control outside reserves. In addition, we demonstrate that there is a trade-off between local and regional conservation objectives when designing reserve networks: small and aggregated reserves based on the extent of dispersal maximize the abundance of corals and herbivores regionally, whereas large and isolated reserves always maximize the abundance of corals within reserves, regardless of the extent of dispersal. The existence of such "conservation traps," which arise from the fulfillment of population-level objectives within local reserves at the cost of community-level objectives at regional scales, suggests the importance of adopting a more holistic strategy to manage complex and interconnected ecosystems.

Bored to Death: Community-Wide Effect of Predation on a Foundation Species in a Low-Disturbance Arctic Subtidal System.

The strength of top-down control by consumers is predicted to decrease with latitude, but most data confirming this assumption come from latitudes <60°, while empirical studies of predation in sub-arctic and arctic marine habitats are few. A barnacle Balanus crenatus is a native foundation species in the shallow subtidal of the White Sea (65° N), hosting a diverse (250+ species) assemblage of macrobenthic organisms. On mixed sediments live barnacles share primary substrates (shells and gravel) with numerous empty barnacle tests, 7% of which had drill holes of an unidentified origin. We manipulated the densities of (i) adult muricid whelks Boreotrophon clathratus (of previously unknown feeding habits), to check if they prey on barnacles, (ii) other predators to reveal their effect on juvenile Boreotrophon, and (iii) empty tests to assess the community-wide effect of predation on barnacles. The abundance of drilled empty tests in the field correlated with that of Boreotrophon. A year-long caging experiment clearly confirmed predation, showing the highest barnacle mortality and proportion of drilled tests in whelk enclosures, and the lowest — in predator exclosure treatments. Boreotrophon preferred the barnacles attached to conspecifics to those from primary substrates. Because of its scarcity Boreotrophon had a minor direct effect on barnacle abundance in the field. Yet, initially defaunated empty tests and live barnacles developed markedly different macrobenthic assemblages, suggesting a strong indirect effect of the predation. Juvenile Boreotrophon were 5-6 times less abundant in open and partial cages than in exclosures and enclosures, which indicates that the recruitment and, consequently, the abundance of Boreotrophon and its predation on Balanus are top-down controlled by apex predators. In contrast, in tropical and temperate intertidal the predation on barnacles is stronger and primarily limited by environmental stress and prey availability.

Does Intraspecific Size Variation in a Predator Affect Its Diet Diversity and Top-Down Control of Prey?

It has long been known that intraspecific variation impacts evolutionary processes, but only recently have its potential ecological effects received much attention. Theoretical models predict that genetic or phenotypic variance within species can alter interspecific interactions, and experiments have shown that genotypic diversity in clonal species can impact a wide range of ecological processes. To extend these studies to quantitative trait variation within populations, we experimentally manipulated the variance in body size of threespine stickleback in enclosures in a natural lake environment. We found that body size of stickleback in the lake is correlated with prey size and (to a lesser extent) composition, and that stickleback can exert top-down control on their benthic prey in enclosures. However, a six-fold contrast in body size variance had no effect on the degree of diet variation among individuals, or on the abundance or composition of benthic or pelagic prey. Interestingly, post-hoc analyses revealed suggestive correlations between the degree of diet variation and the strength of top-down control by stickleback. Our negative results indicate that, unless the correlation between morphology and diet is very strong, ecological variation among individuals may be largely decoupled from morphological variance. Consequently we should be cautious in our interpretation both of theoretical models that assume perfect correlations between morphology and diet, and of empirical studies that use morphological variation as a proxy for resource use diversity.

Evolutionary Trade-Offs in Plants Mediate the Strength of Trophic Cascades

Predators determine herbivore and plant biomass via so-called trophic cascades, and the strength of such effects is influenced by ecosystem productivity. To determine whether evolutionary trade-offs among plant traits influence patterns of trophic control, we manipulated predators and soil fertility and measured impacts of a major herbivore (the aphid Aphis nerii) on 16 milkweed species (Asclepias spp.) in a phylogenetic field experiment. Herbivore density was determined by variation in predation and trade-offs between herbivore resistance and plant growth strategy. Neither herbivore density nor predator effects on herbivores predicted the cascading effects of predators on plant biomass. Instead, cascade strength was strongly and positively associated with milkweed response to soil fertility. Accordingly, contemporary patterns of trophic control are driven by evolutionary convergent trade-offs faced by plants.

Weak top-down control of grazers and periphyton by slimy sculpins in a coldwater stream

Abstract We conducted an enclosure/exclosure experiment during summer 1998 and 1999 to test the strength of top-down control by slimy sculpins (Cottus cognatus) on benthic macroinvertebrates and periphyton. We used 1-m2 cages to manipulate slimy sculpin density within 6 riffles in Valley Creek, Minnesota. Each riffle contained an enclosure stocked with 3 slimy sculpins, a fishless exclosure, and an open control cage. In 1998, densities of total invertebrates and grazers on clay tiles were significantly higher in exclosures than enclosures and control cages; however, densities of individual grazer taxa (Glossosoma and Baetis) on tiles did not differ significantly among treatments. Fish had no significant effect on invertebrates inhabiting gravel substrate. In 1999, densities of total invertebrates and grazers on tiles and inhabiting gravel substrate did not differ significantly among treatments. Slimy sculpins did not have a detectable effect on the drift propensity of invertebrates during either year. Reg...