Predators play an extremely important role in natural communities. In freshwater systems, fish can dominate sorting both at the colonization and post-colonization stage. Specifically, for many colonizing species, fish can have non-lethal, direct effects that exceed the lethal direct effects of predation. Functionally diverse fish species with a range of predatory capabilities have previously been observed to elicit functionally equivalent responses on oviposition in tree frogs. We tested this hypothesis of functional equivalence of non-lethal effects for four predatory fish species, using naturally colonizing populations of aquatic beetles. Among taxa other than mosquitoes, and with the exception of the chemically camouflaged pirate perch, Aphredoderus sayanus, we provide the first evidence of variation in colonization or oviposition responses to different fish species. Focusing on total abundance, Fundulus chrysotus, a gape-limited, surface-feeding fish, elicited unique responses among colonizing Hydrophilidae, with the exception of the smallest and most abundant taxa, Paracymus, while Dytiscidae responded similarly to all avoided fish. Neither family responded to A. sayanus. Analysis of species richness and multivariate characterization of the beetle assemblages for the four fish species and controls revealed additional variation among the three avoided species and confirmed that chemical camouflage in A. sayanus results in assemblages essentially identical to fishless controls. The origin of this variation in beetle responses to different fish is unknown, but may involve variation in cue sensitivity, different behavioral algorithms, or differential responses to species-specific fish cues. The identity of fish species occupying aquatic habitats is crucial to understanding community structure, as varying strengths of lethal and non-lethal effects, as well as their interaction, create complex landscapes of predator effects and challenge the notion of functional equivalence.

Positive correlation of species richness with area is ubiquitous in nature, but the processes driving that relationship, as well as those constraining typical patterns, remain elusive. Patch size variation is pervasive in natural systems, and it is thus critical to understand how variation in patch size, as well as its potential interaction with factors like predation and isolation, affects community assembly. We crossed patch quality (fish presence/absence) with patch size to the examine effects of quality, size, and their interaction on colonization by aquatic insects. Overall, beetles favored small, fishless patches, but individual species sorted across patch size while hemipterans aggregated into large, fishless patches, producing sorting between Coleoptera and Hemiptera. Both patch size and predation risk generated significant variation in community structure and diversity. Patch size preferences for the 14 most abundant species and preeminence of species turnover in patterns of b-diversity reinforce patch size as a driver of regional species sorting via habitat selection. Species sorting at the immigration stage plays a critical role in community assembly. Identifying patch size as a component of perceived quality establishes patch size as a critical niche dimension and alters our view of its role in assembly dynamics and the maintenance of local and regional diversity.

Two of the most important factors determining community structure and diversity within and among habitat patches are patch size and patch quality. Despite the importance of patch size in existing paradigms in island biogeography, metapopulation biology, landscape ecology, and metacommunity ecology, and growing conservation concerns with habitat fragmentation, there has been little investigation into how patch size interacts with patch quality. We crossed three levels of patch size (1.13 m , 2.54 m and 5.73 m ) with two levels of patch quality (fish presence/absence, green sunfish [Lepomis cyanellus] and golden shiners [Notemigonus crysoleucus]) in six replicate experimental landscapes (3 × 2 × 6 = 36 patches). Both fish predators have been previously shown to elicit avoidance in ovipositing treefrogs. We examined how patch size and patch quality, as well as the interaction between size and quality, affected female oviposition preference and male calling site choice in a natural population of treefrogs (Hyla chrysoscelis). Females almost exclusively oviposited in the largest fishless patches, indicating that females use both risk, in the form of fish predators, and size itself, as components of patch quality. Females routinely use much smaller natural and experimental patches, suggesting that the responses to patch size are highly context dependent. Responses to fish were unaffected by patch size. Male responses largely mimicked those of females, but did not drive female oviposition. We suggest that patch size itself functions as another aspect of patch quality for H. chrysoscelis, and serves as another niche dimension across which species may behaviorally sort in natural systems. Because of strong, shared avoidance of fish (as well as other predators), among many colonizing taxa, patch size may be a critical factor in species sorting and processes of community assembly in freshwater habitats, allowing species to behaviorally segregate along gradients of patch size in fishless ponds. Conversely, lack of variation in patch size may concentrate colonization activity, leading to intensification of species interactions and/or increased use of lesser quality patches.

Tree leaf litter inputs to freshwater systems are a major resource and primary drivers of ecosystem processes and structure. Spatial variation in tree species distributions and forest composition control litter inputs across landscapes, but inputs to individual lentic habitat patches are determined by adjacent plant communities. In small, ephemeral, fishless ponds, resource quality and abundance can be the most important factor affecting habitat selection preferences of colonizing animals. We used a landscape of experimental mesocosms to assess how natural populations of aquatic beetles respond over time to variation in tree leaf litter composition (pine or hardwood). Patches with faster-decomposing hardwood leaf litter were initially colonized at higher rates than slower-decomposing pine pools by most species of Hydrophilidae, but this pattern reversed later in the experiment with higher colonization of pine pools by hydrophilids. Colonization did not differ between pine and hardwood for dytiscids and the small hydrophilid Paracymus, but there were distinct beetle assemblages between pine and hardwood patches both early and late in the experiment. Our data support the importance of patch quality and habitat selection as determinants of species abundances, richness, and community structure in freshwater aquatic systems, not only when new habitat patches are formed and initial conditions set, but as patches change due to interactions of processes such as decomposition with time.

Predation risk and resource abundance are two primary characteristics that determine species abundances and community composition. Colonizing organisms should attempt to minimize the risk of mortality and maximize growth through selection of patches with the highest expected fitness. However, maximizing fitness across multiple gradients of patch quality involves accurate cue assessment, integration and behavioural responses that consider multiple factors that affect fitness simultaneously. Our goal was to simultaneously and factorially assess the effects of predation risk and resource abundance among an assemblage of aquatic insects to determine the relative importance of each factor, and whether the two factors interact to affect colonization, oviposition and community assembly. We conducted a field mesocosm experiment in which we crossed predator density (0, 1, 2 fish, Fundulus chrysotus) with supplemental nutrient abundance (0, 4, 8 g rabbit chow) in a 3 × 3 factorial design. We then assayed colonization by natural populations of aquatic beetles and oviposition by natural populations of Culex mosquitoes. We observed species‐specific responses, with many species avoiding fish and some selecting habitats with more nutrients. Nutrients and predator presence only interactively affected oviposition by Culex mosquitoes, and the effect of fish presence exceeded that of nutrients in all but one analysis. Our results illustrate the primacy of predation risk in generating colonization patterns and structuring communities in aquatic habitats, but that colonization responses to variation in multiple components of patch quality are often species‐specific. Simultaneous assessments of multiple aspects of patch quality allow for the determination of potential interactions among cue sources and the relative importance of various patch characteristics to colonizers. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13086/suppinfo is available for this article.