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.