Jarrett E. Byrnes

A global synthesis reveals biodiversity loss as a major driver of ecosystem change

Evidence is mounting that extinctions are altering key processes important to the productivity and sustainability of Earth’s ecosystems. Further species loss will accelerate change in ecosystem processes, but it is unclear how these effects compare to the direct effects of other forms of environmental change that are both driving diversity loss and altering ecosystem function. Here we use a suite of meta-analyses of published data to show that the effects of species loss on productivity and decomposition—two processes important in all ecosystems—are of comparable magnitude to the effects of many other global environmental changes. In experiments, intermediate levels of species loss (21–40%) reduced plant production by 5–10%, comparable to previously documented effects of ultraviolet radiation and climate warming. Higher levels of extinction (41–60%) had effects rivalling those of ozone, acidification, elevated CO2 and nutrient pollution. At intermediate levels, species loss generally had equal or greater effects on decomposition than did elevated CO2 and nitrogen addition. The identity of species lost also had a large effect on changes in productivity and decomposition, generating a wide range of plausible outcomes for extinction. Despite the need for more studies on interactive effects of diversity loss and environmental changes, our analyses clearly show that the ecosystem consequences of local species loss are as quantitatively significant as the direct effects of several global change stressors that have mobilized major international concern and remediation efforts.

The functional role of producer diversity in ecosystems

Over the past several decades, a rapidly expanding field of research known as biodiversity and ecosystem functioning has begun to quantify how the worlds biological diversity can, as an independent variable, control ecological processes that are both essential for, and fundamental to, the functioning of ecosystems. Research in this area has often been justified on grounds that (1) loss of biological diversity ranks among the most pronounced changes to the global environment and that (2) reductions in diversity, and corresponding changes in species composition, could alter important services that ecosystems provide to humanity (e.g., food production, pest/disease control, water purification). Here we review over two decades of experiments that have examined how species richness of primary producers influences the suite of ecological processes that are controlled by plants and algae in terrestrial, marine, and freshwater ecosystems. Using formal meta-analyses, we assess the balance of evidence for eight fundamental questions and corresponding hypotheses about the functional role of producer diversity in ecosystems. These include questions about how primary producer diversity influences the efficiency of resource use and biomass production in ecosystems, how primary producer diversity influences the transfer and recycling of biomass to other trophic groups in a food web, and the number of species and spatial /temporal scales at which diversity effects are most apparent. After summarizing the balance of evidence and stating our own confidence in the conclusions, we outline several new questions that must now be addressed if this field is going to evolve into a predictive science that can help conserve and manage ecological processes in ecosystems.

Invasions and Extinctions Reshape Coastal Marine Food Webs

The biodiversity of ecosystems worldwide is changing because of species loss due to human-caused extinctions and species gain through intentional and accidental introductions. Here we show that the combined effect of these two processes is altering the trophic structure of food webs in coastal marine systems. This is because most extinctions (∼70%) occur at high trophic levels (top predators and other carnivores), while most invasions are by species from lower trophic levels (70% macroplanktivores, deposit feeders, and detritivores). These opposing changes thus alter the shape of marine food webs from a trophic pyramid capped by a diverse array of predators and consumers to a shorter, squatter configuration dominated by filter feeders and scavengers. The consequences of the simultaneous loss of diversity at top trophic levels and gain at lower trophic levels is largely unknown. However, current research suggests that a better understanding of how such simultaneous changes in diversity can impact ecosystem function will be required to manage coastal ecosystems and forecast future changes.

Biodiversity enhances ecosystem multifunctionality across trophic levels and habitats

The importance of biodiversity for the integrated functioning of ecosystems remains unclear because most evidence comes from analyses of biodiversitys effect on individual functions. Here we show that the effects of biodiversity on ecosystem function become more important as more functions are considered. We present the first systematic investigation of biodiversitys effect on ecosystem multifunctionality across multiple taxa, trophic levels and habitats using a comprehensive database of 94 manipulations of species richness. We show that species-rich communities maintained multiple functions at higher levels than depauperate ones. These effects were stronger for herbivore biodiversity than for plant biodiversity, and were remarkably consistent across aquatic and terrestrial habitats. Despite observed tradeoffs, the overall effect of biodiversity on multifunctionality grew stronger as more functions were considered. These results indicate that prior research has underestimated the importance of biodiversity for ecosystem functioning by focusing on individual functions and taxonomic groups.

A meta‐analysis of resource pulse–consumer interactions

Resource pulses are infrequent, large-magnitude, and short-duration events of increased resource availability. They include a diverse set of extreme events in a wide range of ecosystems, but identifying general patterns among the diversity of pulsed resource phenomena in nature remains an important challenge. Here we present a meta-analysis of resource pulse-consumer interactions that addresses four key questions: (1) Which characteristics of pulsed resources best predict their effects on consumers? (2) Which characteristics of consumers best predict their responses to resource pulses? (3) How do the effects of resource pulses differ in different ecosystems? (4) What are the indirect effects of resource pulses in communities? To investigate these questions, we built a data set of diverse pulsed resource-consumer interactions from around the world, developed metrics to compare the effects of resource pulses across disparate systems, and conducted multilevel regression analyses to examine the manner in which variation in the characteristics of resource pulse- consumer interactions affects important aspects of consumer responses. Resource pulse magnitude, resource trophic level, resource pulse duration, ecosystem type and subtype, consumer response mechanisms, and consumer body mass were found to be key explanatory factors predicting the magnitude, duration, and timing of consumer responses. Larger consumers showed more persistent responses to resource pulses, and reproductive responses were more persistent than aggregative responses. Aquatic systems showed shorter temporal lags between peaks of resource availability and consumer response compared to terrestrial systems, and temporal lags were also shorter for smaller consumers compared to larger consumers. The magnitude of consumer responses relative to their resource pulses was generally smaller for the direct consumers of primary resource pulses, compared to consumers at greater trophic distances from the initial resource pulse. In specific systems, this data set showed both attenuating and amplifying indirect effects. We consider the mechanistic processes behind these patterns and their implications for the ecology of resource pulses.

Investigating the relationship between biodiversity and ecosystem multifunctionality: challenges and solutions

Summary Extensive research shows that more species-rich assemblages are generally more productive and efficient in resource use than comparable assemblages with fewer species. But the question of how diversity simultaneously affects the wide variety of ecological functions that ecosystems perform remains relatively understudied. It presents several analytical and empirical challenges that remain unresolved. In particular, researchers have developed several disparate metrics to quantify multifunctionality, each characterizing different aspects of the concept and each with pros and cons. We compare four approaches to characterizing multifunctionality and its dependence on biodiversity, quantifying (i) magnitudes of multiple individual functions separately, (ii) the extent to which different species promote different functions, (iii) the average level of a suite of functions and (iv) the number of functions that simultaneously exceeds a critical threshold. We illustrate each approach using data from the pan-European BIODEPTH experiment and the R multifunc package developed for this purpose, evaluate the strengths and weaknesses of each approach and implement several methodological improvements. We conclude that an extension of the fourth approach that systematically explores all possible threshold values provides the most comprehensive description of multifunctionality to date. We outline this method and recommend its use in future research.

The consequences of consumer diversity loss: different answers from different experimental designs

Predators are often the most vulnerable group to extinction, yet the consequences of changing predator diversity are poorly understood. One source of confusion has been different experimental designs. The multiple-predator effects literature typically employs an additive design, while the biodiversity ecosystem function literature typically uses a replacement design. Separately, these designs each detect only a subset of the changes in food web interactions caused by predator loss. Here, we measure the impact of consumer diversity on sessile marine invertebrates using a combination additive-replacement design. We couple this with a meta-analysis of previous combination experiments. We use these two approaches to explore how each design can detect different types of interactions among predators. We find that, while high diversity does lead to more negative interspecific interactions, the strength of these interactions is often weaker than negative intraspecific interactions caused by increasing the density of a single species alone. We conclude that a hybrid design is the optimal method to explore the mechanisms behind the effects of changing predator diversity. If researchers merely want to know the consequences of changing predator diversity, at a bare minimum, the experimental design must mimic the actual changes in both predator density and diversity in their system of interest. However, only a hybrid design can distinguish the consequences of shifting the balance of interspecific and intraspecific interactions within a community, an issue of great importance when considering both natural diversity loss and pest biocontrol.

Ecological Factors Affecting Community Invasibility

What makes a community invasible? For over a century ecologists have sought to understand the relative importance of biotic and abiotic factors that determine community composition. The fact that we are still exploring this topic today hints at both its importance and complexity. As the impacts from harmful non-native species accumulate, it has become increasingly urgent to find answers to the more applied aspects of this question: what makes a habitat vulnerable to invasion by additional species, and which species are likely to invade? Answers to these questions will not only aid in targeting conservation efforts but will also advance our understanding of marine community ecology. Although the relative importance of abiotic vs. biotic factors in making a habitat invasible varies, abiotic factors undoubtedly serve as the first “filter” to invasions, limiting establishment of non-native (=exotic) species to conditions approximating their native ranges. As obvious examples, tropical corals will not establish in boreal waters, and temperate rocky intertidal species will not colonize tropical shores. Similarly, species cannot invade a community if propagules do not arrive at the site. Other chapters in this volume cover the influence of abiotic factors and propagule supply (Chap. 7, Johnston et al.; Chap. 8, Miller and Ruiz; Chap. 19, Hewitt et al.), so we only briefly review these factors. In this chapter we focus on the question of predicting invasion success of non-native species that are (1) transported to the habitat in question (i.e., propagule supply is not extremely limiting) and (2) physiologically capable of surviving in the climatic regime. We begin with the observation that even in areas of suitable habitat within the current range of an introduced species, there is often dramatic variation in the density, presence, and overall success of the invader. We seek to explain this variation in terms of processes that control the availability of resources. These include not only abiotic and physical factors that determine base resource levels, but also interactions between species or between organisms and their environment that increase resource availability (through disturbance) or decrease resource availability (through competitive processes), or create new resources (through facilitation) (Fig. 12.1).

Short and long term consequences of increases in exotic species richness on water filtration by marine invertebrates

Although recent research has considered the consequences of global declines in the number of species, less attention has focused on the aggregate effects of regional increases in species richness as a result of human-mediated introductions. Here we examine several potential ecosystem consequences of increasing exotic species diversity of suspension feeding marine invertebrates. First, we experimentally manipulated native and non-native suspension feeder richness and measured its effect on short-term phytoplankton clearance rates. Multispecies communities all performed similarly, regardless of whether they were dominated by natives, exotics, or an even mix of the two. Individual species varied considerably in filtration rates, but non-native species often filtered less than the most similar native. Second, we determined potential changes in integrated function over time by comparing seasonal patterns of recruitment as a proxy for the ability to quickly recover filtration capacity after a disturbance. We found that exotic species have complementary seasonal phenologies both to native species and each other. Our results suggest that the consequences of local increases in species richness due to invasions may be manifest over long (annual to interannual) time scales, even when short term changes in ecosystem function are negligible.

Interactions between sea urchin grazing and prey diversity on temperate rocky reef communities

While we frequently observe that increasing species richness within a trophic level can increase the rates of predation or herbivory on lower trophic levels, the general impacts of prey diversity on consumption rates by their predators or herbivores remains unclear. Here we report the results of two field experiments that examined how subcanopy sessile species richness affects rates of consumption by sea urchins. We crossed a natural gradient of species richness in a benthic subtidal community of understory macroalgae and sessile invertebrates against two experimental gradients of urchin density (0-50 and 0-14 individuals) in 0.5-m2 fenced plots. We found that the percent cover of macroalgae and invertebrates consumed by urchins was greater at higher levels of sessile prey species richness. However, this positive association between prey richness and sea urchin consumption was only apparent at low urchin densities; at high urchin densities nearly all algal and invertebrate biomass was consumed irrespective of sessile species richness. The positive relationship between prey richness and urchin consumption was also stronger when the abundance of prey species was more even (i.e., higher Simpsons evenness). Collectively, our results show that the consumptive impacts of urchins on kelp forest understory communities increases as a function of species diversity (both prey richness and evenness), but that prey diversity becomes irrelevant when urchins reach high densities.