Ongoing anthropogenic changes are affecting the abundance, richness, and spatial distribution
of consumers, often leading to regime shifts and the degradation of ecosystem processes (Silliman et
al. 2013). This is especially true for insect herbivores, which influence numerous ecological and
evolutionary processes, including functional attributes relating to plant production and pollination
(Crawley 1989; Branson et al. 2006; Schmitz 2010; Agrawal et al. 2012; Silliman et al. 2013; Borer et
al. 2014). There is widespread evidence for anthropogenic-based changes to insect herbivores, with
signals including reduced species richness, invasion, outbreaks, and selection towards generalist
feeding strategies (Tscharntke & Brandl 2004; Branson et al. 2006; Clavel et al. 2010; Martinson &
Fagan 2014). What remains unclear are the main drivers of these changes, given that human
disturbances may directly affect herbivore diversity and composition (e.g., habitat loss, stand
perturbation) but also do so indirectly via trophically mediated effects on plants and predators
(Halpern et al. 2005; Shurin et al. 2012; Rzanny et al. 2013).
As with all consumer groups, theoretical models for the regulation of insect herbivores
generally emphasize interactions between resource-based bottom-up and predatory top-down
processes (Strong et al. 1984; Crawley 1989; Schmitz 2008; Price et al. 2011). Both processes can be
affected by human activity, but the consequences for herbivore diversity and composition can be
unclear (Silliman et al. 2013). Eutrophication, for example, can elevate production of insect herbivores
by increasing plant productivity (Polis et al. 1997), but simultaneously select against herbivore
richness, especially specialist feeders, because nutrient-rich plant communities tend to be species-poor
(Stevens et al. 2004) and support fewer feeding guilds (Haddad et al. 2000, 2009; Borer et al. 2012).
Loss of apex arthropod predators (Hendrickx et al. 2007; Shochat et al. 2008), however, can
have similar effects, with reduced predation increasing herbivore abundance but also lowering richness if one or a few herbivores dominate (Holt 1977; Oliver et al. 2009). There may also be strong
interactions between human-induced changes to resources and predators: nutrient enrichment may
drive consumer birth rates that far exceed predator-driven herbivore mortality, or cause system
instability by creating top-heavy feeding webs (Polis et al. 1997; McCann 2011; Shurin et al. 2012;
Tunney et al. 2012).
Food web dynamics can also be influenced by anthropogenic landscape transformations through
factors such as habitat loss and patch isolation (Kruess & Tscharntke 1994; Polis et al. 1997;
Valladares et al. 2006). This can affect herbivores spatial turnover directly by dispersal constraints, but
also indirectly by constraining their resources and predators. Habitat loss, for example, can maximize
either stochastic influences on plant community assembly (i.e., different plant communities on
different islands due to random dispersal), or deterministic ones by favoring the same subsets of
species with traits for long-distance colonization (Arroyo-RodrÃguez et al. 2013; Harvey &
MacDougall 2014). Habitat loss can also modify predation pressure by concentrating predators in
smaller remnant areas, creating predation-free patches via dispersal limitation (Kruess & Tscharntke
1994; Hein & Gillooly 2011), or reducing predator diversity on smaller patches because of food
limitation (Holt 1997; Gravel et al. 2011).
Here, I test how multiple stressors on consumers and resources affect insect herbivore diversity,
composition and spatial turnover in grasslands. I use a large-scale factorial mainland-islands food web
assembly experiment, simulating a range of co-occurring changes associated with anthropogenic
impacts (eutrophication, stand perturbation, and habitat fragmentation) and examining their impacts on
the regulation of insect herbivore communities. My main objective is to test whether these typical
anthropogenic impacts associated with global change affect insect herbivores mainly via direct effects
or indirectly by their effects on plant or predator composition and diversity. I meet this objective in three complementary steps: (i) I test for the main effects of habitat size, isolation, eutrophication and
defoliation on herbivore diversity, composition and among-islands spatial turnover, (ii) I use a path
model comparison approach to test whether these effects on herbivores occur through direct or food
web mediated effects, and (iii) I contrast how these changes compare to herbivore dynamics in
unperturbed and continuous mainland habitat. This approach allows me to separate the direct effects of
perturbation on herbivores, versus those mediated indirectly by how perturbations affect plants,
predators, and spatial constraints on herbivores.