Background/Question/Methods Classic trait-based community assembly theory suggests that the simil... more Background/Question/Methods Classic trait-based community assembly theory suggests that the similarity of traits among co-occurring species is shaped by two processes: abiotic filtering and competition. Abiotic filtering, mostly dominating in stressful environments, promotes similarity, while competition, dominating in productive environments, limits similarity. In line with this theory, several studies have found within-community trait similarity to decline along productivity gradients. However, these studies have always focused on species from single trophic levels. Here, we studied how interactions between different trophic levels affect trait similarity patterns along environmental gradients. We proposed three hypotheses for the main drivers of variation in trait similarity patterns of grasses and herbivorous grasshoppers along environmental gradients: 1) environmental control of both, 2) bottom-up control of grasshopper trait similarity, and 3) top-down control of grass trait s...
exclosures contained 30 % taller vegetation than megaherbivore exclosures and they were dominated... more exclosures contained 30 % taller vegetation than megaherbivore exclosures and they were dominated by different grass and grasshopper species. grasshoppers in complete ungulate exclosures were on average 3.5 mm longer than grasshoppers in megaherbivore exclosures, possibly due to changes in plant communities or vegetation structure. We conclude that surprisingly, in this megaherbivore hotspot, mesoherbivores, instead of megaherbivores, most strongly affect grasshopper communities.
Trait-based community assembly theory suggests that trait variation among cooccurring species is ... more Trait-based community assembly theory suggests that trait variation among cooccurring species is shaped by two main processes: abiotic filtering, important in stressful environments and promoting similarity, and competition, more important in productive environments and promoting dissimilarity. Previous studies have indeed found trait similarity to decline along productivity gradients. However, these studies have always been done on single trophic levels. Here, we investigated how interactions between trophic levels affect trait similarity patterns along environmental gradients. We propose three hypotheses for the main drivers of trait similarity patterns of plants and herbivores along environmental gradients: (1) environmental control of both, (2) bottom-up control of herbivore trait variation, and topdown control of grass trait variation.
Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provi... more Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.
There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunc... more There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunctionality), thus ensuring the delivery of ecosystem services important for human well-being. However, the mechanisms underlying this relationship are poorly understood, especially in natural ecosystems. We develop a novel approach to partition biodiversity effects on multifunctionality into three mechanisms and apply this to European forest data. We show that throughout Europe, tree diversity is positively related with multifunctionality when moderate levels of functioning are required, but negatively when very high function levels are desired. For two well-known mechanisms, 'complementarity' and 'selection', we detect only minor effects on multifunctionality. Instead a third, so far overlooked mechanism, the 'jack-of-all-trades' effect, caused by the averaging of individual species effects on function, drives observed patterns. Simulations demonstrate that jack-of-all-trades effects occur whenever species effects on different functions are not perfectly correlated, meaning they may contribute to diversity-multifunctionality relationships in many of the world's ecosystems.
Proceedings of the National Academy of Sciences, 2016
Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to mai... more Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (β-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between β-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.
Background/Question/Methods Recent studies on biodiversity and ecosystem functioning have demonst... more Background/Question/Methods Recent studies on biodiversity and ecosystem functioning have demonstrated the importance of a high diversity for maintaining multiple ecosystem functions simultaneously at high performance (multifunctionality). However, these studies have mostly been carried out with experimental, artificial grassland communities. Evidence whether diversity also promotes multifunctionality in natural forest systems is mostly lacking. Furthermore, it is unknown at which scale diversity has its strongest effect on multifunctionality in natural systems. Here, with the FunDivEurope research consortium, we collected data on tree diversity and multiple ecosystem functions (e.g. productivity, decomposition, pathogen and herbivore resistance, tree regeneration) in 209 forest plots spread over six different countries (from the Mediterranean to the boreal) in Europe. We investigated how both α, β and γ diversity affect multifunctionality. Results/Conclusions We found that abiotic ...
Both arthropods and large grazing herbivores are important components and drivers of biodiversity... more Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: (i) unintentional predation and increased disturbance, (ii) decreases in total resource abundance for arthropods (biomass) and (iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio-temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top-down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity.
Proceedings of the Royal Society B: Biological Sciences, 2009
Predation plays a central role in evolutionary processes, but little is known about how predators... more Predation plays a central role in evolutionary processes, but little is known about how predators affect the expression of heritable variation, restricting our ability to predict evolutionary effects of predation. We reared families of three-spined stickleback Gasterosteus aculeatus from two populations-one with a history of fish predation (predator sympatric) and one without (predator naive)-and experimentally manipulated experience of predators during ontogeny. For a suite of ecologically relevant behavioural ('personality') and morphological traits, we then estimated two key variance components, additive genetic variance (V A ) and residual variance (V R ), that jointly shape narrow-sense heritability (h 2 Z V A /(V A C V R )). Both population and treatment differentially affected V A versus V R , hence h 2 , but only for certain traits. The predator-naive population generally had lower V A and h 2 values than the predator-sympatric population for personality behaviours, but not morphological traits. Values of V R and h 2 were increased for some, but decreased for other personality traits in the predator-exposed treatment. For some personality traits, V A and h 2 values were affected by treatment in the predator-naive population, but not in the predator-sympatric population, implying that the latter harboured less genetic variation for behavioural plasticity.
The grass layer of African savannas consists of two main vegetation types: grazing lawns, dominat... more The grass layer of African savannas consists of two main vegetation types: grazing lawns, dominated by short, mostly clonally reproducing grasses, and bunch grasslands, dominated by tall bunch grasses. This patchy distribution of vegetation types is mostly created by large herbivores, which selectively feed on the more nutritious lawn grass species. Besides grazing, herbivores trample the soil, thereby causing soil compaction, with possible consequences for water infiltration. This raises two questions: (i) is water more limiting in grazing lawns than in bunch grasslands and (ii) are lawn grasses more drought tolerant than bunch grasses? To study these questions, we compared drought conditions in both lawn and bunch grasslands in a South African savanna. Additionally, in a climate room, we compared the performance of three lawn and three bunch grass species under a control and a water limitation treatment. Thirdly, we investigated whether there are differences between lawn and bunch grasses in traits related to drought tolerance. Our results show that despite large differences in water availability in the field, lawn and bunch grasses did not differ in their growth response to drought. Drought reduced growth of both growth forms equally. However, we found strong intrinsic trait differences between growth forms, with lawn grasses having higher specific root length and relative growth rate and bunch grasses having a higher root:shoot ratio. These results suggest that after drought-induced plant death, lawn grasses might be more capable of recolonizing patches of bare soil.
Parasites can affect host phenotypes, influencing their ecology and evolution. Host morphological... more Parasites can affect host phenotypes, influencing their ecology and evolution. Host morphological changes occurring post-infection might result from pathological by-products of infection, or represent adaptations of hosts or parasites. We investigated the morphology of three-spined sticklebacks, Gasterosteus aculeatus, from a population naturally infected with Schistocephalus solidus, which grows to large sizes in their body cavity. We examined local effects of infection on trunk shape, which are imposed directly by the bulk of the growing parasite, and distant effects on head morphology. We show that trunk shape differed between infection classes, and was affected more severely in fish with heavier total parasite mass. We further show unexpected differences in head morphology. The heads of infected fish were reduced in size and differently shaped to those of non-infected fish, with infected fish having deeper heads. Importantly, both head size and shape were also affected more severely in fish with heavier total parasite mass. This latter result suggests that differences in morphology are caused by post-infection changes. Such changes may be incidental, evolutionarily neutral 'side effects' of infection. However, because head morphology affects foraging ecology, such changes are likely to have fitness consequences for hosts, and may constitute adaptations, either of hosts or of parasites. We discuss our finding in the context of the evolution of phenotypic plasticity, and suggest testable hypotheses examining the proximate mechanisms underlying these morphological effects and their potential evolutionary basis.
Background/Question/Methods Classic trait-based community assembly theory suggests that the simil... more Background/Question/Methods Classic trait-based community assembly theory suggests that the similarity of traits among co-occurring species is shaped by two processes: abiotic filtering and competition. Abiotic filtering, mostly dominating in stressful environments, promotes similarity, while competition, dominating in productive environments, limits similarity. In line with this theory, several studies have found within-community trait similarity to decline along productivity gradients. However, these studies have always focused on species from single trophic levels. Here, we studied how interactions between different trophic levels affect trait similarity patterns along environmental gradients. We proposed three hypotheses for the main drivers of variation in trait similarity patterns of grasses and herbivorous grasshoppers along environmental gradients: 1) environmental control of both, 2) bottom-up control of grasshopper trait similarity, and 3) top-down control of grass trait s...
exclosures contained 30 % taller vegetation than megaherbivore exclosures and they were dominated... more exclosures contained 30 % taller vegetation than megaherbivore exclosures and they were dominated by different grass and grasshopper species. grasshoppers in complete ungulate exclosures were on average 3.5 mm longer than grasshoppers in megaherbivore exclosures, possibly due to changes in plant communities or vegetation structure. We conclude that surprisingly, in this megaherbivore hotspot, mesoherbivores, instead of megaherbivores, most strongly affect grasshopper communities.
Trait-based community assembly theory suggests that trait variation among cooccurring species is ... more Trait-based community assembly theory suggests that trait variation among cooccurring species is shaped by two main processes: abiotic filtering, important in stressful environments and promoting similarity, and competition, more important in productive environments and promoting dissimilarity. Previous studies have indeed found trait similarity to decline along productivity gradients. However, these studies have always been done on single trophic levels. Here, we investigated how interactions between trophic levels affect trait similarity patterns along environmental gradients. We propose three hypotheses for the main drivers of trait similarity patterns of plants and herbivores along environmental gradients: (1) environmental control of both, (2) bottom-up control of herbivore trait variation, and topdown control of grass trait variation.
Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provi... more Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.
There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunc... more There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunctionality), thus ensuring the delivery of ecosystem services important for human well-being. However, the mechanisms underlying this relationship are poorly understood, especially in natural ecosystems. We develop a novel approach to partition biodiversity effects on multifunctionality into three mechanisms and apply this to European forest data. We show that throughout Europe, tree diversity is positively related with multifunctionality when moderate levels of functioning are required, but negatively when very high function levels are desired. For two well-known mechanisms, 'complementarity' and 'selection', we detect only minor effects on multifunctionality. Instead a third, so far overlooked mechanism, the 'jack-of-all-trades' effect, caused by the averaging of individual species effects on function, drives observed patterns. Simulations demonstrate that jack-of-all-trades effects occur whenever species effects on different functions are not perfectly correlated, meaning they may contribute to diversity-multifunctionality relationships in many of the world's ecosystems.
Proceedings of the National Academy of Sciences, 2016
Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to mai... more Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (β-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between β-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.
Background/Question/Methods Recent studies on biodiversity and ecosystem functioning have demonst... more Background/Question/Methods Recent studies on biodiversity and ecosystem functioning have demonstrated the importance of a high diversity for maintaining multiple ecosystem functions simultaneously at high performance (multifunctionality). However, these studies have mostly been carried out with experimental, artificial grassland communities. Evidence whether diversity also promotes multifunctionality in natural forest systems is mostly lacking. Furthermore, it is unknown at which scale diversity has its strongest effect on multifunctionality in natural systems. Here, with the FunDivEurope research consortium, we collected data on tree diversity and multiple ecosystem functions (e.g. productivity, decomposition, pathogen and herbivore resistance, tree regeneration) in 209 forest plots spread over six different countries (from the Mediterranean to the boreal) in Europe. We investigated how both α, β and γ diversity affect multifunctionality. Results/Conclusions We found that abiotic ...
Both arthropods and large grazing herbivores are important components and drivers of biodiversity... more Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: (i) unintentional predation and increased disturbance, (ii) decreases in total resource abundance for arthropods (biomass) and (iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio-temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top-down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity.
Proceedings of the Royal Society B: Biological Sciences, 2009
Predation plays a central role in evolutionary processes, but little is known about how predators... more Predation plays a central role in evolutionary processes, but little is known about how predators affect the expression of heritable variation, restricting our ability to predict evolutionary effects of predation. We reared families of three-spined stickleback Gasterosteus aculeatus from two populations-one with a history of fish predation (predator sympatric) and one without (predator naive)-and experimentally manipulated experience of predators during ontogeny. For a suite of ecologically relevant behavioural ('personality') and morphological traits, we then estimated two key variance components, additive genetic variance (V A ) and residual variance (V R ), that jointly shape narrow-sense heritability (h 2 Z V A /(V A C V R )). Both population and treatment differentially affected V A versus V R , hence h 2 , but only for certain traits. The predator-naive population generally had lower V A and h 2 values than the predator-sympatric population for personality behaviours, but not morphological traits. Values of V R and h 2 were increased for some, but decreased for other personality traits in the predator-exposed treatment. For some personality traits, V A and h 2 values were affected by treatment in the predator-naive population, but not in the predator-sympatric population, implying that the latter harboured less genetic variation for behavioural plasticity.
The grass layer of African savannas consists of two main vegetation types: grazing lawns, dominat... more The grass layer of African savannas consists of two main vegetation types: grazing lawns, dominated by short, mostly clonally reproducing grasses, and bunch grasslands, dominated by tall bunch grasses. This patchy distribution of vegetation types is mostly created by large herbivores, which selectively feed on the more nutritious lawn grass species. Besides grazing, herbivores trample the soil, thereby causing soil compaction, with possible consequences for water infiltration. This raises two questions: (i) is water more limiting in grazing lawns than in bunch grasslands and (ii) are lawn grasses more drought tolerant than bunch grasses? To study these questions, we compared drought conditions in both lawn and bunch grasslands in a South African savanna. Additionally, in a climate room, we compared the performance of three lawn and three bunch grass species under a control and a water limitation treatment. Thirdly, we investigated whether there are differences between lawn and bunch grasses in traits related to drought tolerance. Our results show that despite large differences in water availability in the field, lawn and bunch grasses did not differ in their growth response to drought. Drought reduced growth of both growth forms equally. However, we found strong intrinsic trait differences between growth forms, with lawn grasses having higher specific root length and relative growth rate and bunch grasses having a higher root:shoot ratio. These results suggest that after drought-induced plant death, lawn grasses might be more capable of recolonizing patches of bare soil.
Parasites can affect host phenotypes, influencing their ecology and evolution. Host morphological... more Parasites can affect host phenotypes, influencing their ecology and evolution. Host morphological changes occurring post-infection might result from pathological by-products of infection, or represent adaptations of hosts or parasites. We investigated the morphology of three-spined sticklebacks, Gasterosteus aculeatus, from a population naturally infected with Schistocephalus solidus, which grows to large sizes in their body cavity. We examined local effects of infection on trunk shape, which are imposed directly by the bulk of the growing parasite, and distant effects on head morphology. We show that trunk shape differed between infection classes, and was affected more severely in fish with heavier total parasite mass. We further show unexpected differences in head morphology. The heads of infected fish were reduced in size and differently shaped to those of non-infected fish, with infected fish having deeper heads. Importantly, both head size and shape were also affected more severely in fish with heavier total parasite mass. This latter result suggests that differences in morphology are caused by post-infection changes. Such changes may be incidental, evolutionarily neutral 'side effects' of infection. However, because head morphology affects foraging ecology, such changes are likely to have fitness consequences for hosts, and may constitute adaptations, either of hosts or of parasites. We discuss our finding in the context of the evolution of phenotypic plasticity, and suggest testable hypotheses examining the proximate mechanisms underlying these morphological effects and their potential evolutionary basis.
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