Recent work has suggested that emergent ecological network structure exhibits very little spatial or temporal variance despite changes in community composition. However, the changes in network interactions associated with turnover in community composition have seldom been assessed. Here we examine whether changes in ecological networks are best detected by standard emergent network metrics or by assessing internal network changes (i.e. interaction and composition turnover). To eliminate possible spatial or phylogenetic effects, that in large‐scale studies may obscure mechanisms structuring networks and interactions, we sampled multiple antagonistic (plant–herbivore) networks for a single diverse plant family (the Restionaceae) in the hyperdiverse Cape Floristic Region. These are the first plant–herbivore networks constructed for this global biodiversity hotspot. We found invariant emergent network structure despite considerable changes in insect and plant composition across communities over time and space. In contrast, there was high interaction turnover between networks. Seasonally, this was driven by turnover in insect species and insect host switching. Spatially, this was driven by simultaneous turnover in plant and insect species, suggesting that many insects are host specific or that both groups exhibit parallel responses to environmental gradients. Spatial interaction turnover was also driven by turnover in plants, showing that many insects can utilise multiple (possibly closely related) hosts and this may create divergent selection gradients that promote insect speciation. Thus we show highly variable interaction fidelity, despite invariant emergent network structure. We suggest that evaluating internal network changes may be more effective at elucidating the processes structuring networks, and many fine‐scale changes may be obscured when only calculating emergent network metrics.

Summary1. Patterns of niche partitioning can result from local ecological interactions (e.g. interspecific competition) occurring within a contemporary time frame (realized niche partitioning). Alternatively, they may represent the end product of historical processes acting over long time frames (fundamental niche partitioning). 2. Niche partitioning is often detected by analysing patterns of resource use within communities, but experiments are rarely conducted to test whether patterns of non-overlapping resource use reflect realized or fundamental niche partitioning. 3. We studied a community of restio leafhoppers from the genus Cephalelus and their host plants, the Restionaceae (restios). We used network and experimental approaches to determine whether network modularity (a measure of niche partitioning within local communities) reflects fundamental or realized niche partitioning. Using a weighted modularity index for two party networks (e.g. insect-plant), we determined whether the network of this community is modular (i.e. consists of groups of species interacting strongly, with weak interactions between groups). We also aimed to identify specific Cephalelus -restio modules (groups). Using knowledge of module membership to design experiments, we tested whether Cephalelus species from two different modules, Cephalelus uncinatus and Cephalelus pickeri, prefer and perform better on restios from their own modules vs. restios from other modules. These experiments were performed under controlled conditions, eliminating the influences of competition and predation on host choices. 4. The Cephalelus -restio community was modular, implying niche partitioning. Cephalelus also preferred and performed better on restios from their own modules in the absence of local contemporary factors. Most niche partitioning in the investigated Cephalelus community is not caused by local interactions, and thus, host use patterns represent fundamental niches. Our findings highlight the importance of understanding local community structure in the light of processes extrinsic to the local community context.

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BackgroundThe evolution of reproductive isolation between herbivorous insect populations is often initiated by shifts to novel host-plants, a process that underlies some of the best examples of ecological speciation. However, it is not well understood why host-shifts occur. Arguably the most common hypothesis is that host-shifts occur in response to competition, while a less frequently invoked hypothesis is that herbivores adapt locally to geographic differences in potential host-plant communities. Here we investigate whether geographic variation in host-plant availability is likely to have driven host-shifts in restio leafhoppers. We studied local adaptation of a camouflaged restio leafhopper species, Cephalelus uncinatus, to host-plants in the Restionaceae (restios); a family of plants with exceptional diversity in the anomalously species-rich Cape Floristic Region (CFR). To determine whether C. uncinatus experiences heterogeneous host communities across its range, we first quantified the degree of geographic overlap between C. uncinatus and each of its associated host-plant species. Then we quantified trait divergence (host preference, body size and colour) for three pairs of C. uncinatus populations found on different host-plant species differing in their degree of spatial overlap. Spectral reflectance was modelled in bird visual space to investigate whether body colour divergence in C. uncinatus corresponds to leaf sheath colour differences between restio species as perceived by potential predators.ResultsWe demonstrate that C. uncinatus is forced to use different restio species in different regions because of turnover in available host species across its range. Comparisons between geographically separated populations were consistent with local adaptation: restio leafhoppers had preferences for local host-plants over alternative host-plants and matched local plants better in terms of size and colour.ConclusionsSpatial turnover in host-plant availability has likely facilitated host-shifts in C. uncinatus. Spatial turnover in host-plant availability may be an important driver of insect diversification in the CFR and globally.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-017-0916-0) contains supplementary material, which is available to authorized users.