Dirk Sanders

Ecosystem engineering and predation: the multi-trophic impact of two ant species

1. Ants are ubiquitous ecosystem engineers and generalist predators and are able to affect ecological communities via both pathways. They are likely to influence any other terrestrial arthropod group either directly or indirectly caused by their high abundance and territoriality. 2. We studied the impact of two ant species common in Central Europe, Myrmica rubra and Lasius niger, on an arthropod community. Colony presence and density of these two ant species were manipulated in a field experiment from the start of ant activity in spring to late summer. 3. The experiment revealed a positive influence of the presence of one ant colony on densities of decomposers, herbivores and parasitoids. However, in the case of herbivores and parasitoids, this effect was reversed in the presence of two colonies. 4. Generally, effects of the two ant species were similar with the exception of their effect on Braconidae parasitoid densities that responded positively to one colony of M. rubra but not of L. niger. 5. Spider density was not affected by ant colony manipulation, but species richness of spiders responded positively to ant presence. This effect was independent of ant colony density, but where two colonies were present, spider richness was significantly greater in plots with two M. rubra colonies than in plots with one colony of each ant species. 6. To test whether the positive ecosystem engineering effects were purely caused by modified properties of the soil, we added in an additional experiment (i) the soil from ant nests (without ants) or (ii) unmodified soil or (iii) ant nests (including ants) to experimental plots. Ant nest soil on its own did not have a significant impact on densities of decomposers, herbivores or predators, which were significantly, and positively, affected by the addition of an intact nest. 7. The results suggest an important role of both ant species in the grassland food web, strongly affecting the densities of decomposers, herbivores and higher trophic levels. We discuss how the relative impact via bottom-up and top-down effects of ants depends on nest density, with a relatively greater top-down predatory impact at higher densities.

The loss of indirect interactions leads to cascading extinctions of carnivores

Species extinctions are biased towards higher trophic levels, and primary extinctions are often followed by unexpected secondary extinctions. Currently, predictions on the vulnerability of ecological communities to extinction cascades are based on models that focus on bottom-up effects, which cannot capture the effects of extinctions at higher trophic levels. We show, in experimental insect communities, that harvesting of single carnivorous parasitoid species led to a significant increase in extinction rate of other parasitoid species, separated by four trophic links. Harvesting resulted in the release of prey from top-down control, leading to increased interspecific competition at the herbivore trophic level. This resulted in increased extinction rates of non-harvested parasitoid species when their host had become rare relative to other herbivores. The results demonstrate a mechanism for horizontal extinction cascades, and illustrate that altering the relationship between a predator and its prey can cause wide-ranging ripple effects through ecosystems, including unexpected extinctions.

Artificial nighttime light changes aphid-parasitoid population dynamics

Artificial light at night (ALAN) is recognized as a widespread and increasingly important anthropogenic environmental pressure on wild species and their interactions. Understanding of how these impacts translate into changes in population dynamics of communities with multiple trophic levels is, however, severely lacking. In an outdoor mesocosm experiment we tested the effect of ALAN on the population dynamics of a plant-aphid-parasitoid community with one plant species, three aphid species and their specialist parasitoids. The light treatment reduced the abundance of two aphid species by 20% over five generations, most likely as a consequence of bottom-up effects, with reductions in bean plant biomass being observed. For the aphid Megoura viciae this effect was reversed under autumn conditions with the light treatment promoting continuous reproduction through asexuals. All three parasitoid species were negatively affected by the light treatment, through reduced host numbers and we discuss induced possible behavioural changes. These results suggest that, in addition to direct impacts on species behaviour, the impacts of ALAN can cascade through food webs with potentially far reaching effects on the wider ecosystem.

Indirect commensalism promotes persistence of secondary consumer species

Local species extinctions may lead to, often unexpected, secondary extinctions. To predict these, we need to understand how indirect effects, within a network of interacting species, affect the ability of species to persist. It has been hypothesized that the persistence of some predators depends on other predator species that suppress competitively dominant prey to low levels, allowing a greater diversity of prey species, and their predators, to coexist. We show that, in experimental insect communities, the absence of one parasitoid wasp species does indeed lead to the extinction of another that is separated by four trophic links. These results highlight the importance of a holistic systems perspective to biodiversity conservation and the necessity to include indirect population dynamic effects in models for predicting cascading extinctions in networks of interacting species.

Large variation of suction sampling efficiency depending on arthropod groups, species traits, and habitat properties.

Suction sampling is widely used to estimate arthropod abundance and diversity. To test the reliability of abundance data derived from suction sampling, we examined sampling efficiency across a wide range of arthropod groups and tested for effects of species traits, vegetation density, and differences between sites. Suction sampling efficiency was quantified by vacuuming an enclosed meadow area and subsequent removal of the turf, which was treated with heat extraction to collect the remaining arthropods. We obtained 250 pairs of suction and turf samples from seven grasslands with variable vegetation density. High suction sampling efficiencies between 49 and 86% were obtained for Auchenorrhyncha, Heteroptera, Araneida, Curculionoidea, Hymenoptera, and Diptera. In contrast, efficiencies were below 30% for Aphidae, Thysanoptera, Staphylinidae and other Coleoptera, and for soil arthropods such as Collembola, Isopoda, Diplopoda, and Formicidae. Efficiency varied significantly among habitats (sites) for most groups, often more than two‐fold. Surprisingly, sampling efficiency for Hymenoptera, Diplopoda, and Collembola increased with vegetation density, probably because aboveground activity of these taxa was higher in dense vegetation. Suction sampling was nearly twice as efficient for spiders living in the vegetation than for spiders living near the soil surface, and cursorial and large‐bodied spider species were more efficiently sampled than web‐builders and small species. Depending on the sampling effort, suction sampling missed between 49% (one sample) and 31% (250 samples) of the spider species present. Suction sampling efficiency varied more strongly among sites and among arthropod groups than previously recognized. Abundance data derived from suction sampling are strongly underestimated, especially for arthropods living near the soil surface. Thus, comparisons of abundance and diversity between sites should be restricted to vegetation‐dwelling species of the most efficiently sampled groups. The positive relationship of sampling efficiency with vegetation density demonstrates that variation in efficiency is mediated by arthropod behaviour.

Defensive insect symbiont leads to cascading extinctions and community collapse

Abstract Animals often engage in mutualistic associations with microorganisms that protect them from predation, parasitism or pathogen infection. Studies of these interactions in insects have mostly focussed on the direct effects of symbiont infection on natural enemies without studying community‐wide effects. Here, we explore the effect of a defensive symbiont on population dynamics and species extinctions in an experimental community composed of three aphid species and their associated specialist parasitoids. We found that introducing a bacterial symbiont with a protective (but not a non‐protective) phenotype into one aphid species led to it being able to escape from its natural enemy and increase in density. This changed the relative density of the three aphid species which resulted in the extinction of the two other parasitoid species. Our results show that defensive symbionts can cause extinction cascades in experimental communities and so may play a significant role in the stability of consumer‐herbivore communities in the field.

Individual and species‐specific traits explain niche size and functional role in spiders as generalist predators

The function of a predator within a community is greatly based on its trophic niche, that is the number and the strength of feeding links. In generalist predators, which feed on a wide range of prey, the size and position of the trophic niche is likely determined by traits such as hunting mode, the stratum they occur in, their body size and age. We used stable isotope analyses ((13)C and (15)N) to measure the trophic niche size of nine spider species within a forest hedge community and tested for species traits and individual traits that influence stable isotope enrichment, niche size and resource use. The spiders Enoplognatha, Philodromus, Floronia, and Heliophanus had large isotopic niches, which correspond to a more generalistic feeding behaviour. In contrast, Araneus, Metellina and Agelena, as top predators in the system, had rather narrow niches. We found a negative correlation between trophic position and niche size. Differences in trophic position in spiders were explained by body size, hunting modes and stratum, while niche size was influenced by hunting mode. In Philodromus, the size of the trophic niche increased significantly with age. Fitting spiders to functional groups according to their mean body size, hunting mode and their habitat domain resulted in largely separated niches, which indicates that these traits are meaningful for separating functional entities in spiders. Functional groups based on habitat domain (stratum) caught the essential functional differences between the species with species higher up in the vegetation feeding on flying insects and herb and ground species also preying on forest floor decomposers. Interestingly, we found a gradient from large species using a higher habitat domain and having a smaller niche to smaller species foraging closer to the ground and having a larger niche. This shows that even within generalist predators, such as spiders, there is a gradient of specialism that can be predicted by functional traits.

The impact of an ant–aphid mutualism on the functional composition of the secondary parasitoid community

1. Mutualistic and antagonistic interactions, although often studied independently, may affect each other, and food web dynamics are likely to be determined by the two processes working in concert.

Experimental Evidence for the Population-Dynamic Mechanisms Underlying Extinction Cascades of Carnivores

Species extinction rates due to human activities are high, and initial extinctions can trigger cascades of secondary extinctions, leading to further erosion of biodiversity. A potential major mechanism for secondary extinction cascades is provided by the long-standing theory that the diversity of consumer species is maintained due to the positive indirect effects that these species have on each other by reducing competition among their respective resource species. This means that the loss of one carnivore species could lead to competitive exclusion at the prey trophic level, leading to extinctions of further carnivore species. Evidence for these effects is difficult to obtain due to many confounding factors in natural systems, but extinction cascades that could be due to this mechanism have been demonstrated in simplified laboratory microcosms. We established complex insect food webs in replicated field mesocosms and found that the overharvesting of one parasitoid wasp species caused increased extinction rates of other parasitoid species, compared to controls, but only when we manipulated the spatial distribution of herbivore species such that the potential for interspecific competition at this level was high. This provides clear evidence for horizontal extinction cascades at high trophic levels due to the proposed mechanism. Our results demonstrate that the loss of carnivores can have widespread effects on other species at the same trophic level due to indirect population-dynamic effects that are rarely considered in this context.

Effects of urbanization on direct and indirect interactions in a tri-trophic system.

While effects of urbanization on species assemblages are receiving increasing attention, effects on ecological interactions remain largely unexplored. We investigated how urbanization influences the strength of direct and indirect trophic interactions in a tri- trophic system. In a field experiment including five cities and nearby farmed areas, we used potted Vicia faba plants and manipulated the presence of Megoura viciae aphids and that of naturally occurring aphid predators. When predators could access aphids, they reduced their abundance less in the urban than in the agricultural ecosystem. Compared to aphid abundance on plants without predator access, abundance on plants with predator access was 2.58 times lower in urban and 5.27 times lower in agricultural areas. This indicates that urbanization limited top-down control of aphids by predators. In both ecosystems, plant biomass was negatively affected by herbivores and positively affected by predators, but the positive indirect predator effect was weaker in cities. Compared to aphid-infested plants without predator access, plants with predator access were 1.89 times heavier in urban and 2.12 times heavier in agricultural areas. Surprisingly, differences between ecosystems regarding the indirect predator effect on plants were not explained by the differentially strong herbivore suppression. Instead, the urban environment limited plant biomass per se, thereby mitigating the scope of a positive predator effect. Our results show that urbanization can influence direct and indirect trophic interactions through effects on biotic top-down forces and on plant growth. In order to understand how urbanization affects biodiversity and ecosystem functioning, it is fundamental to not only consider species assemblages, but also species interactions.