Food web structure shaped by habitat size and climate across a latitudinal gradient.

Habitat size and climate are known to affect the trophic structure and dynamics of communities, but their interactive effects are poorly understood. Organisms from different trophic levels vary in terms of metabolic requirements and heat dissipation. Indeed, larger species such as keystone predators require more stable climatic conditions than their prey. Likewise, habitat size disproportionally affects large-sized predators, which require larger home ranges and are thus restricted to larger habitats. Therefore, food web structure in patchy ecosystems is expected to be shaped by habitat size and climate variations. Here we investigate this prediction using natural aquatic microcosm (bromeliad phytotelmata) food webs composed of litter resources (mainly detritus), detritivores, mesopredators, and top predators (damselflies). We surveyed 240 bromeliads of varying sizes (water retention capacity) across 12 open restingas in SE Brazil spread across a wide range of tropical latitudes (-12.6° to -27.6°, ca. 2,000 km) and climates (Δ mean annual temperature = 5.3°C). We found a strong increase in predator-to-detritivore mass ratio with habitat size, which was representative of a typical inverted trophic pyramid in larger ecosystems. However, this relationship was contingent among the restingas; slopes of linear models were steeper in more stable and favorable climates, leading to inverted trophic pyramids (and top-down control) being more pronounced in environments with more favorable climatic conditions. By contrast, detritivore-resource and mesopredator-detritivore mass ratios were not affected by habitat size or climate variations across latitudes. Our results highlight that the combined effects of habitat size, climate and predator composition are pivotal to understanding the impacts of multiple environmental factors on food web structure and dynamics.

Predation by odonates depresses mosquito abundance in water-filled tree holes in Panama.

In the lowland moist forest of Barro Colorado Island (BCI), Panama, larvae of four common species of odonates, a mosquito, and a tadpole are the major predators in water-filled tree holes. Mosquito larvae are their most common prey. Holes colonized naturally by predators and prey had lower densities of mosquitoes if odonates were present than if they were absent. Using artificial tree holes placed in the field, we tested the effects of odonates on their mosquito prey while controlling for the quantity and species of predator, hole volume, and nutrient input. In large and small holes with low nutrient input, odonates depressed the number of mosquitoes present and the number that survived to pupation. Increasing nutrient input (and consequently, mosquito abundance) to abnormally high levels dampened the effect of predation when odonates were relatively small. However, the predators grew faster with higher nutrients, and large larvae in all three genera reduced the number of mosquitoes surviving to pupation, even though the abundance of mosquito larvae remained high. Size-selective predation by the odonates is a likely explanation for this result; large mosquito larvae were less abundant in the predator treatment than in the controls. Because species assemblages were similar between natural and artificial tree holes, our results suggest that odonates are keystone species in tree holes on BCI, where they are the most common large predators.

Community organization in streams: the importance of species interactions, physical factors, and chance.

Experimental studies were used to examine the mechanisms governing the distribution and abundance of two major patch types in unshaded reaches of Augusta Creek, Michigan (USA). One patch type is dominated by Cladophora glomerata, a macroalga potentially able to monopolize space, whereas the other type is comprised of a low-growing, epilithic microalgal lawn inhabited by several species of sessile grazers (especially the caddisflies Leucotrichia pictipes and Psychomyia flavida). Cladophora patches are absent from mid-channel sites characterized by current velocities ≤ ca. 50 cm s-1; caging experiments indicate that their absence is due to grazing by crayfish (Orconectes propinquus). Cladophora's presence in sites with velocities >50 cm s-1 apparently results in part because crayfish foraging activity is impaired in high flow regimes. The presence of Cladophora strongly affects various other invertebrates due to its alteration of abiotic and biotic characteristics of the microhabitat. For example, the abundance of sessile grazers (e.g. Leucotrichia and Psychomyia) that inhabit microalgal patches is negatively correlated to the abundance of Cladophora, whereas the abundance of several other invertebrates (e.g. Stenonema mayflies and Taeniopteryx stoneflies) is positively correlated to Cladophora's abundance. Therefore, in some portions of this system, crayfish act as keystone predators because of their ability to regulate the abundance of Cladophora, which in turn has strong positive and negative effects on other components of the community. Cladophora does not always monopolize space at high velocities in the absence of crayfish, however. If sessile grazers arrive at such sites before Cladophora, they can prevent its establishment. Thus, where crayfish are absent, the likelihood that a site will be dominated by either Cladophora patches or sessile grazer - microalgal lawn patches depends on two sets of stochastic processes: (1) those that create bare space (e.g. disturbance and grazer emergence); and (2) those controlling the timing of recruitment by Cladophora or grazers at these bare sites. These priority effects (i.e. the ability of grazers and Cladophora to inhibit each other's establishment) contribute to the marked spatial heterogeneity of these two patch types. Collectively, these results demonstrate how interactions between competition, predation, and physical factors can generate a complex mixture of community patterns.