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.     

Effects of trophic skewing of species richness on ecosystem functioning in a diverse marine community

Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.     

Mass–abundance scaling in avian communities is maintained after tropical selective logging

1. Selective logging dominates forested landscapes across the tropics. Despite the structural damage incurred, selectively logged forests typically retain more biodiversity than other forest disturbances. Most logging impact studies consider conventional metrics, like species richness, but these can conceal subtle biodiversity impacts. The mass–abundance relationship is an integral feature of ecological communities, describing the negative relationship between body mass and population abundance, where, in a system without anthropogenic influence, larger species are less abundant due to higher energy requirements. Changes in this relationship can indicate community structure and function changes.
2. We investigated the impacts of selective logging on the mass–abundance scaling of avian communities by conducting a meta‐analysis to examine its pantropical trend. We divide our analysis between studies using mist netting, sampling the understory avian community, and point counts, sampling the entire community.
3. Across 19 mist‐netting studies, we found no consistent effects of selective logging on mass–abundance scaling relative to primary forests, except for the omnivore guild where there were fewer larger‐bodied species after logging. In eleven point‐count studies, we found a more negative relationship in the whole community after logging, likely driven by the frugivore guild, showing a similar pattern.
4. Limited effects of logging on mass–abundance scaling may suggest high species turnover in logged communities, with like‐for‐like replacement of lost species with similar‐sized species. The increased negative mass–abundance relationship found in some logged communities could result from resource depletion, density compensation, or increased hunting; potentially indicating downstream impacts on ecosystem functions.
5. Synthesis and applications. Our results suggest that size distributions of avian communities in logged forests are relatively robust to disturbance, potentially maintaining ecosystem processes in these forests, thus underscoring the high conservation value of logged tropical forests, indicating an urgent need to focus on their protection from further degradation and deforestation.

     

Top predator removals have consistent effects on large species despite high environmental variability

Top predator losses affect a wide array of ecological processes, and there is growing evidence that top predators are disproportionately vulnerable to environmental changes. Despite increasing recognition of the fundamental role that top predators play in structuring communities and ecosystems, it remains challenging to predict the consequences of predator extinctions in highly variable environments. Both biotic and abiotic drivers determine community structure, and manipulative experiments are necessary to disentangle the effects of predator loss from other co‐occurring environmental changes. To explore the consistency of top predator effects in ecological communities that experience high local environmental variability, we experimentally removed top predators from arid‐land stream pool mesocosms in southeastern Arizona, USA, and measured natural background environmental conditions. We inoculated mesocosms with aquatic invertebrates from local streams, removed the top predator Abedus herberti (Hemiptera: Belostomatidae) from half of the mesocosms as a treatment, and measured community divergence at the end of the summer dry season. We repeated the experiment in two consecutive years, which represented two very different biotic and abiotic environments. We found that some of the effects of top predator removal were consistent despite significant differences in environmental conditions, community composition, and colonist sources between years. As in other studies, top predator removal did not affect overall species richness or abundance in either year, and we observed inconsistent effects on community and trophic structure. However, top predator removal consistently affected large‐bodied species (those in the top 1% of the community body size distribution) in both years, increasing the abundance of mesopredators and decreasing the abundance of detritivores, even though the identity of these species varied between years. Our findings highlight the vulnerability of large taxa to top predator extirpations and suggest that the consistency of observed ecological patterns may be as important as their magnitude.     

Ecological correlates of body size in gamasid mites parasitic on small mammals: abundance and niche breadth

We studied ecological correlates of body size (abundance and niche breadth) in gamasid mites parasitic on small mammals in 28 regions of the Palearctic. We predicted that smaller species would be characterized by higher abundance than larger species, all else (e.g. host species) being equal. We also predicted that host specificity of mites would decrease (that is, number of host species they use would increase) with an increase in their body size. We focused on mites collected from host bodies that include a) species that feed solely on host’s blood (obligate exclusive haematophages), b) species that feed on both host’s blood and small arthropods (obligate non‐exclusive haematophages), and c) facultative haematophages. We expected that the relationship between body size and abundance and/or host specificity would be more pronounced in obligate exclusively haematophagous mites than for obligate non‐exclusively and facultative haematophagous mites. Across all mite species across regions, mean abundance correlated negatively with body size. The same was true for obligate haematophagous species, but not for facultative haematophages. When mite communities on the same host in a location were considered, the negative body mass–abundance relationship was found in only 3 of 44 communities. Nevertheless, a meta‐analytic (across host species) estimate of the slope of this relationship appeared to be significantly negative. No significant relationship between mite body size and host specificity was found in the analyses across all mite species as well as in obligate exclusive or obligate non‐exclusive haematophages. However, the number of hosts used by facultative haematophagous mites decreased significantly with an increase in their body size. We explain the relationships between morphological (body size) and ecological (abundance and niche breadth) properties of ectoparasites by their interactions with hosts or physical environment.     

Size‐balanced community reorganization in response to nutrients and warming

It is widely accepted that global warming will adversely affect ecological communities. As ecosystems are simultaneously exposed to other anthropogenic influences, it is important to address the effects of climate change in the context of many stressors. Nutrient enrichment might offset some of the energy demands that warming can exert on organisms by stimulating growth at the base of the food web. It is important to know whether indirect effects of warming will be as ecologically significant as direct physiological effects. Declining body size is increasingly viewed as a universal response to warming, with the potential to alter trophic interactions. To address these issues, we used an outdoor array of marine mesocosms to examine the impacts of warming, nutrient enrichment and altered top‐predator body size on a community comprised of the predator (shore crab Carcinus maenas), various grazing detritivores (amphipods) and algal resources. Warming increased mortality rates of crabs, but had no effect on their moulting rates. Nutrient enrichment and warming had near diametrically opposed effects on the assemblage, confirming that the ecological effects of these two stressors can cancel each other out. This suggests that nutrient‐enriched systems might act as an energy refuge to populations of species under metabolic constraints due to warming. While there was a strong difference in assemblages between mesocosms containing crabs compared to mesocosms without crabs, decreasing crab size had no detectable effect on the amphipod or algal assemblages. This suggests that in allometrically balanced communities, the expected long‐term effect of warming (declining body size) is not of similar ecological consequence to the direct physiological effects of warming, at least not over the six week duration of the experiment described here. More research is needed to determine the long‐term effects of declining body size on the bioenergetic balance of natural communities.     

Trophic consequences of a biological invasion: do plant invasions increase predator abundance?

As invasive species become integrated into existing communities, they engage in a wide variety of trophic interactions with other community members. Many of these interactions are direct (e.g. predator–prey interactions or interference competition), but invasive species also can affect native community members indirectly, by influencing the abundances of intermediary species in trophic webs. Observational studies suggest that invasive plant species affect herbivorous arthropod communities and that these effects may flow up trophic webs to influence the abundance of predators. However, few studies have experimentally manipulated the presence of invasive plants to quantify the effects of plant invasion on higher trophic levels. Here, I use comparisons across sites that have or have not been invaded by the invasive plant Medicago polymorpha, combined with experimental removals of Medicago and insect herbivores, to investigate how a plant invasion affects the abundance of predators. Both manipulative and observational experiments showed that Medicago increased the abundance of the exotic herbivore Hypera and predatory spiders, suggesting positive bottom–up effects of plant invasions on higher trophic levels. Path analyses conducted on data from natural habitats revealed that Medicago primarily increased spider abundance through herbivore‐mediated indirect pathways. Specifically, Medicago density was positively correlated with the abundance of the dominant herbivore Hypera, and increased Hypera densities were correlated with increased spider abundance. Smaller‐scale experimental studies confirmed that Medicago may increase spider abundance through herbivore‐mediated indirect pathways, but also showed that the effects of Medicago varied across sites, including having no effect or having direct effects on spider abundance. If effects of invasive species commonly flow through trophic webs, then invasive species have the potential to affect numerous species throughout the community, especially those species whose dynamics are tightly connected to highly‐impacted community members through trophic linkages.     

Non‐consumptive effects of a top‐predator decrease the strength of the trophic cascade in a four‐level terrestrial food web

The fear of predators can strongly impact food web dynamics and ecosystem functioning through effects on herbivores morphology, physiology or behaviour. While non‐consumptive predator effects have been mostly studied in three‐level food chains, we lack evidence for the propagation of non‐consumptive indirect effects of apex predators in four level food‐webs, notably in terrestrial ecosystems. In experimental mesocosms, we manipulated a four‐level food chain including top‐predator cues (snakes), mesopredators (lizards), herbivores (crickets), and primary producers (plants). The strength of the trophic cascade induced by mesopredators through the consumption of herbivores decreased in the presence of top‐predator cues. Specifically, primary production was higher in mesocosms where mesopredators were present relative to mesocosms with herbivores only, and this difference was reduced in presence of top‐predator cues, probably through a trait‐mediated effect on lizard foraging. Our study demonstrates that non‐consumptive effects of predation risk can cascade down to affect both herbivores and plants in a four‐level terrestrial food chain and emphasises the need to quantify the importance of such indirect effects in natural communities.     

Robustness of size–structure across ecological networks in pelagic systems

The study of biomass size distributions has become an important tool for addressing aquatic ecosystem complexity and the consequences of anthropogenic disturbances. However, it remains unclear how changes in pelagic food web topology affect the biomass size–structure. Employing a dynamic multispecies bioenergetic consumer-resource model, we simulated biomass trajectories over time in 10,000 virtual networks of varying topology to address which food web properties are important in determining size–structure in pelagic systems. The slopes of the normalized biomass size spectra (NBSS) and Pareto’s shape parameter (γ) of our modeled communities are consistent with theoretically expected values for steady-state systems and empirical values reported for several aquatic ecosystems. We found that the main drivers of the NBSS slope and Pareto’s γ were the slope of the relationship between body mass and trophic level, the maximum trophic level of the food web, and the stability of total community biomass. Our analyses showed a clear conservative trend in pelagic community size–structure as demonstrated by the robustness of the NBSS slope and Pareto’s γ against most of the topological changes in virtual networks. Nevertheless, these analyses also caution that major disturbances in large-bodied or top-trophic level individuals may disrupt this stable pattern.     

A food web perspective on large herbivore community limitation

The exceptional diversity of large mammals in African savannas provides an ideal opportunity to explore the relative importance of top‐down and bottom‐up controls of large terrestrial herbivore communities. Recent work has emphasized the role of herbivore and carnivore body size in shaping these trophic relationships. However, the lack of across‐ecosystem comparisons using a common methodology prohibits general conclusions. Here we used published data on primary production, herbivore and carnivore densities and diets to estimate the consumption fluxes between three trophic levels in four African savanna ecosystems. Our food web approach suggests that the body size distribution within and across trophic levels has a strong influence on the strength of top‐down control of herbivores by carnivores and on consumption fluxes within ecosystems, as predicted by theoretical food web models. We generalize findings from the Serengeti ecosystem that suggest herbivore species below 150 kg are more likely to be limited by predation. We also emphasize the key functional role played by the largest species at each trophic level. The abundance of the largest herbivore species largely governs the consumption of primary production in resident communities. Similarly, predator guilds in which the largest carnivore species represent a larger share of carnivore biomass are likely to exert a stronger top‐down impact on herbivores. Our study shows how a food web approach allows integrating current knowledge and offers a powerful framework to better understand the functioning of ecosystems.     

Size diversity and species diversity relationships in fish assemblages of Western Palearctic lakes

Body size, coupled with abundance and taxonomy, may help to understand the mechanisms shaping community structure. Since the body size of fish is closely related to their trophic niche, size diversity (based on individual body size) of fish communities may capture intraspecific variations in fish trophic niches that are not detected by species diversity. Thus, the relationship between size diversity and species diversity may help to integrate variation at both intraspecific and interspecific levels. We studied the relationship between species diversity and size diversity as a measure of the degree of overlap in size among species and thereby the potential overlap in niches in a community. We hypothesized that the relationship between size diversity and species would be different across the European continent due to different levels of size overlap in fish communities. The data were derived from samplings of fish communities using standardised benthic gill nets in 363 lakes. At the continental scale, size diversity increased with species diversity; at the ecoregion scale, the slope of the relation changed across the continent, with the greatest mismatch occurring in northern Europe where communities comprised only one or a few species, but each of which exhibited a great range in size. There was an increase in slope towards the south with significant relations for four out of six ecoregions. The steeper size diversity‐species diversity slope at lower latitudes is attributable to a lower overlap in fish size and thus likely to finer niche separation. Our results also suggest that size diversity is not a strong surrogate for species diversity in European lake fish communities. Thus, particularly in fish communities composed of few species, measuring size diversity may help to detect potential functional variation which may be neglected by measuring species diversity alone.     

Characterizing the trophic niches of stocked and resident cyprinid fishes: consistency in partitioning over time,space and body sizes

Hatchery‐reared fish are commonly stocked into freshwaters to enhance recreational angling. As these fishes are often of high trophic position and attain relatively large sizes, they potentially interact with functionally similar resident fishes and modify food‐web structure. Hatchery‐reared barbel Barbus barbus are frequently stocked to enhance riverine cyprinid fish communities in Europe; these fish can survive for over 20 years and exceed 8 kg. Here, their trophic consequences for resident fish communities were tested using cohabitation studies, mainly involving chub Squalius cephalus, a similarly large‐bodied, omnivorous and long‐lived species. These studies were completed over three spatial scales: pond mesocosms, two streams and three lowland rivers, and used stable isotope analysis. Experiments in mesocosms over 100 days revealed rapid formation of dietary specializations and discrete trophic niches in juvenile B. barbus and S. cephalus. This niche partitioning between the species was also apparent in the streams over 2 years. In the lowland rivers, where fish were mature individuals within established populations, this pattern was also generally apparent in fishes of much larger body sizes. Thus, the stocking of these hatchery‐reared fish only incurred minor consequences for the trophic ecology of resident fish, with strong patterns of trophic niche partitioning and diet specialization. Application of these results to decision‐making frameworks should enable managers to make objective decisions on whether cyprinid fish should be stocked into lowland rivers according to ecological risk.     

Predicting the consequences of species loss using size‐structured biodiversity approaches

Understanding the consequences of species loss in complex ecological communities is one of the great challenges in current biodiversity research. For a long time, this topic has been addressed by traditional biodiversity experiments. Most of these approaches treat species as trait‐free, taxonomic units characterizing communities only by species number without accounting for species traits. However, extinctions do not occur at random as there is a clear correlation between extinction risk and species traits. In this review, we assume that large species will be most threatened by extinction and use novel allometric and size‐spectrum concepts that include body mass as a primary species trait at the levels of populations and individuals, respectively, to re‐assess three classic debates on the relationships between biodiversity and (i) food‐web structural complexity, (ii) community dynamic stability, and (iii) ecosystem functioning. Contrasting current expectations, size‐structured approaches suggest that the loss of large species, that typically exploit most resource species, may lead to future food webs that are less interwoven and more structured by chains of interactions and compartments. The disruption of natural body‐mass distributions maintaining food‐web stability may trigger avalanches of secondary extinctions and strong trophic cascades with expected knock‐on effects on the functionality of the ecosystems. Therefore, we argue that it is crucial to take into account body size as a species trait when analysing the consequences of biodiversity loss for natural ecosystems. Applying size‐structured approaches provides an integrative ecological concept that enables a better understanding of each species' unique role across communities and the causes and consequences of biodiversity loss.     

Fish diversity as a determinant of ecosystem properties across multiple trophic levels

Biodiversity has been established as a potential determinant of function in many ecosystems; however, previous research has mostly focused on primary producers and effects at a single trophic level. A broader perspective that considers multiple components of food webs is necessary to understand natural systems. In particular, consumer diversity needs to be more thoroughly examined as trophic interactions and indirect effects can alter ecosystem properties. We test the potential for consumer diversity (fish richness and composition) to govern food web dynamics at two levels of environmental complexity (mesocosms and experimental ponds) and explore the consequences of removing individual species of fish on lower trophic levels. In mesocosms, both the richness and density of zooplankton were reduced when more fish species were present. No effects from the fish treatments were found on phytoplankton, but phosphorus levels increased with higher fish richness. Removing either generalist or specialist fish species increased the richness and density of zooplankton and the amount of phytoplankton, whereas all fish species had redundant effects on nutrients. In ponds, a dominant fish species (specialist shiner) determined the richness and density of zooplankton. In contrast, phytoplankton and nutrients were reduced by higher fish richness in the fall and spring. Overall, the specialist shiner had unique effects on the pond food web suggesting the key to understanding function is the presence of a dominant species and their biological interactions. Differences between mesocosms and ponds are likely due to increased heterogeneity of resources in the ponds allowing species to specialize on different prey. Our study links the biodiversity ecosystem function paradigm with food web concepts to improve predictions for conservation and management actions in response to changes in biodiversity.     

Biodiversity and ecosystem functioning in food webs: the vertical diversity hypothesis

One challenge in merging community and ecosystem ecology is to integrate the complexity of natural multitrophic communities into concepts of ecosystem functioning. Here, we combine food‐web and allometry theories to demonstrate that primary production, as measured by the total nutrient uptake of the multitrophic community, is determined by vertical diversity (i.e. food web's maximum trophic level) and structure (i.e. distributions of species and their abundances and metabolic rates across trophic levels). In natural ecosystems, the community size distribution determines all these vertical patterns and thus the total nutrient uptake. Our model suggests a vertical diversity hypothesis (VDH) for ecosystem functioning in complex food webs. It predicts that, under a given nutrient supply, the total nutrient uptake increases exponentially with the maximum trophic level in the food web and it increases with its maximum body size according to a power law. The VDH highlights the effect of top–down regulation on plant nutrient uptake, which complements traditional paradigms that emphasised the bottom–up effect of nutrient supply on vertical diversity. We conclude that the VDH contributes to a synthetic framework for understanding the relationship between vertical diversity and ecosystem functioning in food webs and predicting the impacts of global changes on multitrophic ecosystems.     

On the determinants of census area: implications for mammalian macroecological patterns

The aim of this study was to examine the effects of various biological factors such as body mass, trophic level, climate and geography on census area in terrestrial mammals. We also examine the effects of census area on the population density–body mass relationship. The geographic areas covered in this study include most major terrestrial biomes including taïga, desert, savanna, grassland, tropical dry forest, temperate dry forest, tropical rain forest and temperate rain forest. An extensive literature search was conducted and we compiled data on census area and body mass from 377 mammalian populations and 59 communities. Statistical analyses include linear regression, Kruskal–Wallis analysis of variance, LOWESS, and multiple regression. Overall, the regression between log census area (A) and log body mass (M) yielded a slope of 0.710, which did not differ significantly from 0.75, but it was significantly different from 1.0. The analyses also showed that the log A–log M relationship is constrained within a well‐defined space in a similar fashion to the home range–body mass relationship. When mammals were separated into trophic groups, regression lines differed significantly in their intercepts, but not in slopes. At the community level, the census area was particularly affected by the population with the largest body mass within the community. Both the number of species and number of taxa encompassed by the community were found to be correlated positively with log A (r = 0.26, P = 0.0464 and r = 0.27, P = 0.0398, respectively). Sampling of mammalian species is not usually random. Not only is census area significantly associated with the technique used to sample a given species, but it is also influenced by biological factors that have been shown previously to influence population density. Striking similarities were found between the census area–body mass relationship and the home range–body mass relationship, suggesting that investigators may sample mammalian populations over areas that actually reflect the use of space of their focal species.     

Exploring predictions of abundance from body mass using hierarchical comparative approaches

Understanding and predicting how and why abundance varies is one of the central questions in ecology. One of the few consistent predictors of variation in abundance between species has been body mass, but the nature of this relationship has been contentious. Here I explore the relationship between body mass and abundance in birds of North America, using hierarchical partitioning of variance and regressions at taxonomic levels above the species. These analyses show that much variation in abundance is found across space, while a moderate amount of variation is found at the species/genus and also at the family/order level. However, body size and trophic level primarily vary at the family/order level, suggesting that mechanisms based on body size and energy should primarily explain only this moderate-sized, taxonomically conserved component of variation in abundance. Body size does explain more than 50% of the variation at this level (and almost 75% when trophic level is also included). This tighter relationship makes clear that energetic equivalence (slope = -3/4) sets an upper limit but does not describe the relationship between body mass and average abundance for birds of North America. Finally, I suggest that this hierarchical, multivariate approach should be used more often in macroecology.     

Abundance–body mass relationships in size-structured food webs

In communities sharing a common energy source, the energetic equivalence hypothesis predicts that numerical abundance (N) scales with body mass (M) as M^?0.75. However, in size‐structured food webs all individuals do not share a common energy source, and the energy available (E) to larger individuals is constrained by inefficient energy transfer through the food chains that support them. This is expected to lead to steeper scalings of N with M. Here, we formalize and test an existing model for predicting abundance–body mass scaling, where the decline in E with M is calculated from the mean predator–prey body mass ratio (from size‐based nitrogen stable isotope analysis) and trophic transfer efficiency. We show that the steep predicted scalings of abundance and body mass (N scales as M^?1.2, B scales as M^?0.2) in a marine food web are consistent with empirical estimates and can be attributed to the small predator–prey body mass ratio (106 : 1). As a previous study has shown that environmental stability may favour low predator–prey mass ratios and long food chains, we predict that steeper abundance–body mass relationships will be found in more stable environments.