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Here, we aim to understand how multiple mechanisms interact within and between life stages to determine frequency‐dependent population growth, which has a key role stabilizing local competitor coexistence.

We conducted field experiments in three lakes manipulating relative frequencies of twoEnallagmadamselfly species to evaluate demographic contributions of three mechanisms affecting different fitness components across the life cycle: the effect of resource competition on individual growth rate, predation shaping mortality rates, and mating harassment determining fecundity. We then used a demographic model that incorporates carry‐over effects between life stages to decompose the relative effect of each fitness component generating frequency‐dependent population growth.

This decomposition showed that fitness components combined to increase population growth rates for one species when rare, but they combined to decrease population growth rates for the other species when rare, leading to predicted exclusion in most lakes.

Because interactions between fitness components within and between life stages vary among populations, these results show that local coexistence is population specific. Moreover, we show that multiple mechanisms do not necessarily increase competitor coexistence, as they can also combine to yield exclusion. Identifying coexistence mechanisms in other systems will require greater focus on determining contributions of different fitness components across the life cycle shaping competitor coexistence in a way that captures the potential for population‐level variation.

We first evaluated how host identity and density could shape parasitism. To test the effects of con‐ and heterospecific host density on parasitism, we used a field experiment withEnallagma basidensandE. signatum. We found that parasitism did not vary with con‐ or heterospecific density and was determined by host identity alone, with no spillover effects.

We also evaluated the potential role of local adaptation and resource availability in shaping parasitism. To do so, we usedE. signatumin a reciprocal transplant experiment crossed with a prey resource‐level manipulation. This experiment revealed that parasitism declined sharply for one host population in its non‐local lake, but not the other source population, with no effects of prey levels. This asymmetry implies that damselflies express enhanced defences against parasitism that are neither population‐specific nor dependent on resource abundance, or that mites developed heightened local host specificity.

The results of multivariate modeling from an observational study generally supported these experimental findings: neither host density nor resource abundance strongly explained among‐population variation in parasitism. Instead, local abiotic conditions (pH) had the strongest relationship with parasitism, with minimal associations with predator density, temperature and a measure of immune function.

Collectively, our findings suggest a crucial role for the local environment in shaping host–parasite interactions within multi‐host–parasite systems. More generally, these results show that research at the intersection of community ecology and disease ecology is critical for understanding host–parasite dynamics within natural communities.

What demographic outcomes of the interactions within and between species in these guilds are expected to produce ecological drift versus coexistence remains unclear. To address this issue, and guide empirical testing, we consider models of a guild of ecologically equivalent competitors feeding on a single resource to explore when community drift should manifest.

We show that community drift dynamics only emerge when the density‐dependent effects of each species on itself are identical to its density‐dependent effects on every other guild member. In contrast, if each guild member directly limits itself more than it limits the abundance of other guild members, all species in the guild are coexisting, even though they all are ecologically equivalent with respect to their interactions with species outside the guild (i.e. resources, predators, mutualists). Hence, considering only interspecific ecological differences generating density dependence, and not fully accounting for the preponderance of mechanisms causing intraspecific density dependence, will provide an incomplete picture for segregating between neutrality and coexistence. We also identify critical experiments necessary to disentangle guilds of ecologically equivalent species from those experiencing ecological drift, as well as provide an overview of ways of incorporating a mechanistic basis into studies of species coexistence and neutrality.

Identifying these characteristics, and the mechanistic basis underlying community structure, is not merely an exercise in clarifying the semantics of coexistence and neutral theories, but rather reflects key differences that must exist among community members in order to determine how and why communities are structured.