Consequences of biodiversity loss for litter decomposition across biomes.

The decomposition of dead organic matter is a major determinant of carbon and nutrient cycling in ecosystems, and of carbon fluxes between the biosphere and the atmosphere. Decomposition is driven by a vast diversity of organisms that are structured in complex food webs. Identifying the mechanisms underlying the effects of biodiversity on decomposition is critical given the rapid loss of species worldwide and the effects of this loss on human well-being. Yet despite comprehensive syntheses of studies on how biodiversity affects litter decomposition, key questions remain, including when, where and how biodiversity has a role and whether general patterns and mechanisms occur across ecosystems and different functional types of organism. Here, in field experiments across five terrestrial and aquatic locations, ranging from the subarctic to the tropics, we show that reducing the functional diversity of decomposer organisms and plant litter types slowed the cycling of litter carbon and nitrogen. Moreover, we found evidence of nitrogen transfer from the litter of nitrogen-fixing plants to that of rapidly decomposing plants, but not between other plant functional types, highlighting that specific interactions in litter mixtures control carbon and nitrogen cycling during decomposition. The emergence of this general mechanism and the coherence of patterns across contrasting terrestrial and aquatic ecosystems suggest that biodiversity loss has consistent consequences for litter decomposition and the cycling of major elements on broad spatial scales.

When does diversity matter? Species functional diversity and ecosystem functioning across habitats and seasons in a field experiment.

Despite ample experimental evidence indicating that biodiversity might be an important driver of ecosystem processes, its role in the functioning of real ecosystems remains unclear. In particular, the understanding of which aspects of biodiversity are most important for ecosystem functioning, their importance relative to other biotic and abiotic drivers, and the circumstances under which biodiversity is most likely to influence functioning in nature, is limited. We conducted a field study that focussed on a guild of insect detritivores in streams, in which we quantified variation in the process of leaf decomposition across two habitats (riffles and pools) and two seasons (autumn and spring). The study was conducted in six streams, and the same locations were sampled in the two seasons. With the aid of structural equations modelling, we assessed spatiotemporal variation in the roles of three key biotic drivers in this process: functional diversity, quantified based on a species trait matrix, consumer density and biomass. Our models also accounted for variability related to different litter resources, and other sources of biotic and abiotic variability among streams. All three of our focal biotic drivers influenced leaf decomposition, but none was important in all habitats and seasons. Functional diversity had contrasting effects on decomposition between habitats and seasons. A positive relationship was observed in pool habitats in spring, associated with high trait dispersion, whereas a negative relationship was observed in riffle habitats during autumn. Our results demonstrate that functional biodiversity can be as significant for functioning in natural ecosystems as other important biotic drivers. In particular, variation in the role of functional diversity between seasons highlights the importance of fluctuations in the relative abundances of traits for ecosystem process rates in real ecosystems.

Continental-scale effects of nutrient pollution on stream ecosystem functioning.

Excessive nutrient loading is a major threat to aquatic ecosystems worldwide that leads to profound changes in aquatic biodiversity and biogeochemical processes. Systematic quantitative assessment of functional ecosystem measures for river networks is, however, lacking, especially at continental scales. Here, we narrow this gap by means of a pan-European field experiment on a fundamental ecosystem process--leaf-litter breakdown--in 100 streams across a greater than 1000-fold nutrient gradient. Dramatically slowed breakdown at both extremes of the gradient indicated strong nutrient limitation in unaffected systems, potential for strong stimulation in moderately altered systems, and inhibition in highly polluted streams. This large-scale response pattern emphasizes the need to complement established structural approaches (such as water chemistry, hydrogeomorphology, and biological diversity metrics) with functional measures (such as litter-breakdown rate, whole-system metabolism, and nutrient spiraling) for assessing ecosystem health.

Placing biodiversity and ecosystem functioning in context: environmental perturbations and the effects of species richness in a stream field experiment.

Greater biodiversity is often associated with increased ecosystem process rates, and is expected to enhance the stability of ecosystem functioning under abiotic stress. However, these relationships might themselves be altered by environmental factors, complicating prediction of the effects of species loss in ecosystems subjected to abiotic stress. In boreal streams, we investigated effects of biodiversity and two abiotic perturbations on three related indices of ecosystem functioning: leaf decomposition, detritivore leaf processing efficiency (LPE) and detritivore growth. Replicate field enclosures containing leaves and detritivore assemblages were exposed to liming and nutrient enrichment, raising pH and nutrient levels. Both treatments constitute perturbations for our naturally acidic and nutrient-poor streams. We also varied detritivore species richness and density. The effects of the abiotic and diversity manipulations were similar in magnitude, but whereas leaf decomposition increased by 18% and 8% following liming and nutrient enrichment, respectively, increased detritivore richness reduced leaf decomposition (6%), detritivore LPE (19%) and detritivore growth (12%). The detritivore richness effect on growth was associated with negative trait-independent complementarity, indicating interspecific interference competition. These interactions were apparently alleviated in both enriched and limed enclosures, as trait-independent complementarity became less negative. LPE increased with detritivore density in the monocultures, indicating benefits of intra-specific aggregation that outweighed the costs of intra-specific competition, and dilution of these benefits probably contributed to lowered leaf decomposition in the species mixtures. Finally, the effects of liming were reduced in most species mixtures relative to the monocultures. These results demonstrate how environmental changes might regulate the consequences of species loss for functioning in anthropogenically perturbed ecosystems, and highlight potential influences of biodiversity on functional stability. Additionally, the negative effects of richness and positive effects of density in our field study were opposite to previous laboratory observations, further illustrating the importance of environmental context for biodiversity-ecosystem functioning relationships.

Contrasting effects of anthropogenic and natural acidity in streams: a meta-analysis.

Large-scale human activities including the extensive combustion of fossil fuels have caused acidification of freshwater systems on a continental scale, resulting in reduced species diversity and, in some instances, impaired ecological functioning. In regions where acidity is natural, however, species diversity and functioning seem to be less affected. This contrasting response is likely to have more than one explanation including the possibility of adaptation in organisms exposed to natural acidity over evolutionary time scales and differential toxicity due to dissimilarities in water chemistry other than pH. However, empirical evidence supporting these hypotheses is equivocal. Partly, this is because previous research has mainly been conducted at relatively small geographical scales, and information on ecological functioning in this context is generally scarce. Our goal was to test whether anthropogenic acidity has stronger negative effects on species diversity and ecological functioning than natural acidity. Using a meta-analytic approach based on 60 datasets, we show that macroinvertebrate species richness and the decomposition of leaf litter -- an important process in small streams -- tend to decrease with increasing acidity across regions and across both the acidity categories. Macroinvertebrate species richness, however, declines three times more rapidly with increasing acidity where it is anthropogenic than where it is natural, in agreement with the adaptation hypothesis and the hypothesis of differences in water chemistry. By contrast, the loss in ecological functioning differs little between the categories, probably because increases in the biomass of taxa remaining at low pH compensate for losses in functionality that would otherwise accompany losses of taxa from acidic systems. This example from freshwater acidification illustrates how natural and anthropogenic stressors can differ markedly in their effects on species diversity and one aspect of ecological functioning.

Diverging effects of anthropogenic acidification and natural acidity on community structure in Swedish streams.

Anthropogenic acidification caused by aerial deposition of acidifying substances is known to have detrimental effects on freshwater biota, including reductions in species diversity and ecosystem functioning. However, such impairment is not found in systems acidified to a similar extent by natural processes. A proposed explanation for this difference is that freshwater organisms have had far more time to evolve and adapt to natural than anthropogenic acidification. Thus, where acidity is natural, adaptation may account for diverse and functional communities. Here, we investigated whether adaptations--that were previously implied to occur on small spatial scales--may explain the species richness patterns on a much larger geographical scale, apply to ecological functioning, and are relevant in Sweden, where natural acidity is geologically relatively recent. Therefore, we compared differences in species diversity and ecosystem process rates between 24 acidic and circumneutral streams in northern Sweden, where acidity is natural, and southern Sweden, where acidity is largely anthropogenic. In agreement with our predictions, the difference in macroinvertebrate species richness between acidic and circumneutral streams was threefold larger in the region where acidity was anthropogenic than where it was natural, albeit marginally non-significantly. In contrast, no such trend was found for the rates of decomposition by microbes and leaf-feeding macroinvertebrates, possibly due to functional redundancy. The structure of species assemblages differed between acidic and circumneutral sites and between the regions. Our results agree with the notion that freshwater biota are adapted to natural acidity, but competing explanations including other differences in water chemistry and differences in the biogeographical colonization histories may also account for part of the observed patterns. Since naturally acidic environments similar to those in northern Sweden are widespread, we predict that diverse and functionally efficient freshwater communities that are well adapted to such conditions are more common than currently recognized.

Vertebrate host specificity of wild-caught blackflies revealed by mitochondrial DNA in blood.

Blood-feeding blackflies (Diptera: Simuliidae) transmit pathogens, harass vertebrate hosts and may cause lethal injuries in attacked victims, but with traditional methods it has proved difficult to identify their hosts. By matching mitochondrial DNA (mtDNA) sequences in blood collected from engorged blackflies with stored sequences in the GenBank database, relationships between 17 blackfly species and 25 species of vertebrate hosts were revealed. Our results demonstrate a predominance of large hosts and marked discrimination between blackflies using either avian or mammalian hosts. Such information is of vital interest in studies of disease transmission, coevolutionary relationships, population ecology and wildlife management.

Testing hypotheses on egg number and size in black flies (Diptera: Simuliidae).

Field-collected gravid females were used to test a series of inter- and intraspecific hypotheses on egg size and number in 23 species of black flies. Qualitative differences between bird and mammal blood did not result in significant differences with respect to fecundity properties. The five species with overwintering larvae produced more eggs than did the 18 species with overwintering eggs. Contrary to prediction, egg size was correlated significantly with body size in only two of 17 examined species; indeed, in three species this relationship was negative. However, the association between female size and egg number within species was positive. Strong support was found for a tradeoff between egg size and number across species but not within species. The present study shows similarities with other dipterans but also substantial variability, probably reflecting a high degree of flexibility. The results are primarily discussed in an evolutionary context, but they also provide useful information for black fly management.

Mechanisms behind positive diversity effects on ecosystem functioning: testing the facilitation and interference hypotheses.

Little is known about the mechanisms behind positive effects of species richness on ecosystem functioning. In a previous study that showed a positive effect of aquatic detritivore species richness on leaf litter breakdown (process) rates, we proposed that facilitation and release from intra-specific interference were the two most likely mechanisms. To test the interference hypothesis, we performed an experiment using three densities of each of three detritivore species and found varying effects on leaf breakdown rates across species: one species showed no effect, one a positive, marginally insignificant, effect, and a third species showed a significant, positive effect of decreasing density. The density (interference) effect thus partly explained the results from our previous study. The facilitation hypothesis was tested by sequentially introducing and removing two species. We predicted that, if this hypothesis were true, facilitation would be expressed in higher process rates than when replacing with individuals of the same species. We found that process rate per unit biomass did increase when one species was introduced after the other species, while the opposite sequence did not show any increase. Hence, this result was also confirmative of our previous results. Therefore, we conclude that both intra-specific interference and inter-specific facilitation may explain the positive effect of species richness observed in our system. Since many species exhibit intra-specific interference that inhibits foraging efficiency, this may be a general mechanism generating effects of species richness per se. If facilitation is unidirectional, or if it involves few species, it is more likely to be species specific with species identities being more important than species richness per se. We conclude that species loss may be expected to have negative consequences on ecosystem functioning if anyspecies is lost, with additional effects in the event of losing "facilitator" species.

Simulating species loss following perturbation: assessing the effects on process rates.

We removed stream-living macroinvertebrate shredder species in the sequences in which they are predicted to disappear, in response to two common types of anthropogenic disturbances: acidification and organic pollution, and analysed the effects on leaf breakdown rates. The experiment was performed in field microcosms using three shredder species. Species identity significantly affected leaf breakdown rates, while species richness per se was non-significant. The simulated sequential species loss showed large effects on leaf breakdown rates, with observed rates being significantly higher than expected from single-species treatments in two, out of four, two-species, and in all four three-species treatments. The invertebrates used in this study were taxonomically distinct (Insecta: Plecoptera and Trichoptera; Crustacea: Amphipoda), and of different sizes, hence a high degree of complementarity was probably present. A method to study the effects of species loss, characteristic of perturbation type, could be more useful than a random approach when investigating the impact of perturbation. Our results may have general applicability for investigations on the effects of diversity loss on ecosystem functioning in any ecosystem exposed to human perturbations, given that the order of extinction is known or can easily be assessed.

Changing risk of predation for a filter-feeding insect along a current velocity gradient.

Flume experiments were carried out to examine whether larval blackflies (Simulium ornatum complex, Diptera: Simuliidae) use microhabitats with a highvelocity current to reduce the risk of predation by some of their main predators, viz. larvae of the stoneflies Isoperla grammatica and Diura nanseni (Plecoptera: Perlodidae), and the caddis-fly Rhyacophila nubila (Trichoptera: Rhyacophilidae). We exposed blackfly larvae to four different current velocities and measured their feeding rate using dye particles. The maximum feeding rate was recorded at intermediate velocities (18.8 and 36.2 cm/s), whereas at low (7.3 cm/s) and high (53.3 cm/s) velocities, the feeding rate was reduced. In separate experiments, we investigated the behaviour and attack success of the different predator species. The two perlodids showed a similar hunting behaviour, which was significantly less successful at higher velocities. Drift of the perlodids from the experimental arena resulted in reduced encounter and attack rates, especially in I. grammatica, which had completely lost efficiency at 36.2 cm/s. R. nubila had a slower mode of hunting and was unaffected by current speed within the velocity gradient studied. Drift in Rhyacophila was rare. Observations on the behaviour of blackfly larvae were performed in the same experiments. The larvae showed no apparent ability to sense the presence of the predators except when these disrupted the flow pattern or were in physical contact, which often resulted in aggressive defence, though without effect on the predators. Escape of blackfly larvae by drift did occur, but this was no more common than being captured. In a current velocity gradient, blackfly larvae showed a weak preference for increasing velocities. Thus, at velocities between 7 and 54 cm/s, blackfly larvae appear to select microhabitats with high current velocities, despite a reduction in feeding optimality, thereby easing the predation impact from perlodids, though not from Rhyacophila. The study demonstrates the importance of microhabitat selection by blackfly larvae both for efficient feeding and predator avoidance.

Interactions between the leech Glossiphonia complanata and its gastropod prey.

Predator-prey interactions between the predatory leech, Glossiphonia complanata, and its gastropod prey were investigated in laboratory experiments, including behavioural observations with the aid of time-lapse video technique. Six gastropod species were investigated, viz. Lymnaea peregra, Planorbis planorbis, Physa fontinalis, Ancylus fluviatilis, Bithynia tentaculata, and Theodoxus fluviatilis. The species studied exhibited anti-predator defences, which had their maximum efficiency at different stages of encounter with G. complanata. The activity of B. tentaculata decreased with increasing leech activity, but was relatively higher when food was present than when not. Handling times were dependent on the time elapsed since the previous meal was captured (intercatch interval), which in turn was related to the size of the previous prey. Handling time was also related to the size of both predator and prey. The capture efficiency was high for small prey and the leeches spent more time in patches with higher yield. They were, however, unable to discriminate between patches of different prey density.