Diane S. Srivastava

Biodiversity loss and its impact on humanity

The most unique feature of Earth is the existence of life, and the most extraordinary feature of life is its diversity. Approximately 9 million types of plants, animals, protists and fungi inhabit the Earth. So, too, do 7 billion people. Two decades ago, at the first Earth Summit, the vast majority of the world’s nations declared that human actions were dismantling the Earth’s ecosystems, eliminating genes, species and biological traits at an alarming rate. This observation led to the question of how such loss of biological diversity will alter the functioning of ecosystems and their ability to provide society with the goods and services needed to prosper.

Effects of biodiversity on the functioning of trophic groups and ecosystems

Over the past decade, accelerating rates of species extinction have prompted an increasing number of studies to reduce species diversity experimentally and examine how this alters the efficiency by which communities capture resources and convert those into biomass. So far, the generality of patterns and processes observed in individual studies have been the subjects of considerable debate. Here we present a formal meta-analysis of studies that have experimentally manipulated species diversity to examine how it affects the functioning of numerous trophic groups in multiple types of ecosystem. We show that the average effect of decreasing species richness is to decrease the abundance or biomass of the focal trophic group, leading to less complete depletion of resources used by that group. At the same time, analyses reveal that the standing stock of, and resource depletion by, the most species-rich polyculture tends to be no different from that of the single most productive species used in an experiment. Of the known mechanisms that might explain these trends, results are most consistent with what is called the ‘sampling effect’, which occurs when diverse communities are more likely to contain and become dominated by the most productive species. Whether this mechanism is widespread in natural communities is currently controversial. Patterns we report are remarkably consistent for four different trophic groups (producers, herbivores, detritivores and predators) and two major ecosystem types (aquatic and terrestrial). Collectively, our analyses suggest that the average species loss does indeed affect the functioning of a wide variety of organisms and ecosystems, but the magnitude of these effects is ultimately determined by the identity of species that are going extinct.

Impacts of plant diversity on biomass production increase through time because of species complementarity

Accelerating rates of species extinction have prompted a growing number of researchers to manipulate the richness of various groups of organisms and examine how this aspect of diversity impacts ecological processes that control the functioning of ecosystems. We summarize the results of 44 experiments that have manipulated the richness of plants to examine how plant diversity affects the production of biomass. We show that mixtures of species produce an average of 1.7 times more biomass than species monocultures and are more productive than the average monoculture in 79% of all experiments. However, in only 12% of all experiments do diverse polycultures achieve greater biomass than their single most productive species. Previously, a positive net effect of diversity that is no greater than the most productive species has been interpreted as evidence for selection effects, which occur when diversity maximizes the chance that highly productive species will be included in and ultimately dominate the biomass of polycultures. Contrary to this, we show that although productive species do indeed contribute to diversity effects, these contributions are equaled or exceeded by species complementarity, where biomass is augmented by biological processes that involve multiple species. Importantly, both the net effect of diversity and the probability of polycultures being more productive than their most productive species increases through time, because the magnitude of complementarity increases as experiments are run longer. Our results suggest that experiments to date have, if anything, underestimated the impacts of species extinction on the productivity of ecosystems.

Why more productive sites have more species: an experimental test of theory using tree-hole communities.

One of the most common explanations for an increase in species richness with productivity is what we have dubbed the “More Individuals Hypothesis.” According to this hypothesis, more productive sites can support higher total abundances and, since species richness is an increasing function of total abundance, so will it be of productivity. This hypothesis assumes that communities are limited by productivity. We tested the More Individuals Hypothesis using the detritivorous aquatic insect communities of tree holes. When tree holes with varying levels of productivity (debris amount) were allowed to be colonized (through oviposition), more productive tree holes did have more species but not more individuals. Neither was total energy use strictly proportional to productivity. Only in communities forced to disassemble through productivity reductions were the predictions of the More Individuals Hypothesis satisfied. Ovipositing adults may prefer productive tree holes not because they contain more resources but because they are anticipated to be less likely to dry out. In tree holes, and more generally, the More Individuals Hypothesis is an insufficient explanation for increases in species richness with productivity because it neither accounts for the different processes of local coloni zation and extinction nor allows body size to correlate with extinction risk.

Phylogenetic diversity and the functioning of ecosystems

Phylogenetic diversity (PD) describes the total amount of phylogenetic distance among species in a community. Although there has been substantial research on the factors that determine community PD, exploration of the consequences of PD for ecosystem functioning is just beginning. We argue that PD may be useful in predicting ecosystem functions in a range of communities, from single-trophic to complex networks. Many traits show a phylogenetic signal, suggesting that PD can estimate the functional trait space of a community, and thus ecosystem functioning. Phylogeny also determines interactions among species, and so could help predict how extinctions cascade through ecological networks and thus impact ecosystem functions. Although the initial evidence available suggests patterns consistent with these predictions, we caution that the utility of PD depends critically on the strength of phylogenetic signals to both traits and interactions. We advocate for a synthetic approach that incorporates a deeper understanding of how traits and interactions are shaped by evolution, and outline key areas for future research. If these complexities can be incorporated into future studies, relationships between PD and ecosystem function bear promise in conceptually unifying evolutionary biology with ecosystem ecology.

A POSITIVE FEEDBACK : HERBIVORY, PLANT GROWTH, SALINITY, AND THE DESERTIFICATION OF AN ARCTIC SALT-MARSH

1 A 2-year study is described which suggests that a positive feedback process results in the destruction of salt-marsh swards and the exposure of bare sediments at La Perouse Bay, Manitoba, Canada. Lesser snow geese initiate the process by grubbing for roots and rhizomes of salt-marsh graminoids (Puccinellia phryganodes and Carex subspathacea) in spring. The increased rates of evaporation from sediments beneath disturbed or destroyed swards in summer result in high soil salinities that adversely affect the growth of the remaining grazed plants. 2 Above-ground biomass and soil salinity differed between sites in the salt marsh. Soil salinity was inversely related to above-ground biomass and shoot density of Puccinellia phryganodes. Increased biomass led to reduced soil salinity at sites where exclosures were erected. 3 Plant growth, measured as the rate of leaf births on Puccinellia shoots, was reduced by high soil salinities at sites where exclosures were erected. 4 Leaf demography of transplanted experimental plants of Puccinellia differed in 1992, but not 1991, between plants transplanted into sites with different amounts of above-ground biomass. Leaf births and deaths were highest for plants grown in sites where above-ground biomass was high and lowest for plants transplanted into bare sites. Grazing had no effect on leaf demography in 1991 and only marginally increased the rate of leaf deaths in 1992. 5 Growth of transplanted individuals of Carex subspathacea was similarly highest at sites where the standing crop of Puccinellia and Carex was high and was lowest in bare sites. 6 Algal crusts, which formed on bare or poorly vegetated sites, also reduced the growth of Puccinellia plants. 7 The effects of this deleterious positive feedback on plant growth are discussed in relation to changes occurring in the lesser snow goose colonies at La Perouse Bay and elsewhere.

THE LOCAL–REGIONAL RELATIONSHIP: IMMIGRATION, EXTINCTION, AND SCALE

While local processes (e.g., competition, predation, and disturbance) pre- sumably cause species exclusion and thus limit diversity in individual communities, regional processes (e.g., historical events, immigration, and speciation) are assumed to provide a source of species to colonize and thus enrich local communities. Ecologists have attempted to distinguish between these two sets of processes using graphical evidence for local as- semblage saturation. However, such efforts have been controversial and are antithetical to the fact that local diversity bears an imprint of both. We examine the local-regional species richness relationship from the perspective of the theory of island biogeography and develop a model that can produce the full range of observed local-regional richness relationships from linear to curvilinear. Importantly, unlike previous models, we do not require species interactions to produce the curvilinear pattern. Curvilinear relationships arise if per-species stochastic extinction rates are substantially higher than colonization rates, while linear relationships result if colonization rates are higher than extinction rates. Because we also show that merely changing the sampling scale can make local-regional relationships appear either saturated or unsaturated, an inference of ecological processes, derived solely from local-regional relationships, is unwarranted.

Habitat structure, trophic structure and ecosystem function: interactive effects in a bromeliad-insect community.

Although previous studies have shown that ecosystem functions are affected by either trophic structure or habitat structure, there has been little consideration of their combined effects. Such interactions may be particularly important in systems where habitat and trophic structure covary. I use the aquatic insects in bromeliads to examine the combined effects of trophic structure and habitat structure on a key ecosystem function: detrital processing. In Costa Rican bromeliads, trophic structure naturally covaries with both habitat complexity and habitat size, precluding any observational analysis of interactions between factors. I therefore designed mesocosms that allowed each factor to be manipulated separately. Increases in mesocosm complexity reduced predator (damselfly larva) efficiency, resulting in high detritivore abundances, indirectly increasing detrital processing rates. However, increased complexity also directly reduced the per capita foraging efficiency of the detritivores. Over short time periods, these trends effectively cancelled each other out in terms of detrital processing. Over longer time periods, more complex patterns emerged. Increases in mesocosm size also reduced both predator efficiency and detritivore efficiency, leading to no net effect on detrital processing. In many systems, ecosystem functions may be impacted by strong interactions between trophic structure and habitat structure, cautioning against examining either effect in isolation.

Reducing horizontal and vertical diversity in a foodweb triggers extinctions and impacts functions

Species loss can result in secondary extinctions and changes in ecosystem functions at distant trophic levels. Such effects of species loss are predicted to be affected by both the number of species lost within a trophic level (horizontal diversity) and the number of trophic levels lost (vertical diversity). We experimentally manipulated horizontal and vertical diversity within an aquatic insect community, and examined responses throughout the food web. Horizontal and vertical diversity both impacted ciliates: reduction of detritivorous insect diversity resulted in secondary extinctions and decreased density of ciliates, but only when an insect predator was simultaneously absent. Horizontal and vertical diversity differed in their effect on other foodweb processes, including detrital processing, predator growth, and densities of rotifers, flagellates and flatworms. These results caution that foodweb effects of multitrophic species loss may not be reliably predicted from manipulations of just one dimension of diversity.

Why Are Predators More Sensitive to Habitat Size than Their Prey? Insights from Bromeliad Insect Food Webs

Ecologists have hypothesized that the exponent of species‐area power functions (z value) should increase with trophic level. The main explanation for this pattern has been that specialist predators require prior colonization of a patch by their prey, resulting in a compounding of the effects of area up trophic levels. We propose two novel explanations, neither of which assumes trophic coupling between species. First, sampling effects can result in different z values if the abundances of species differ (in mean or evenness) between trophic levels. Second, when body size increases between trophic levels, effects of body size on z values may appear as differences between trophic levels. We test these alternative explanations using invertebrate food webs in 280 bromeliads from three countries. The z value of predators was higher than that of prey. Much of the difference in z values could be explained by sampling effects but not by body size effects. When damselflies occurred in the species pool, predator z values were even higher than predicted, as damselflies avoid small, drought‐prone bromeliads. In one habitat, dwarf forests, detrital biomass became decoupled from bromeliad size, which also caused large trophic differences in z values. We argue that there are often simpler explanations than trophic coupling to explain differences in z values between trophic levels.