Half a million species of herbivorous insects have been described. Most of them are diet specialists, using only a few plant species as hosts. Biologists suspect that their specificity is key to their diversity. But why do herbivorous insects tend to be diet specialists? In this review, we catalog a broad range of explanations. We review the evidence for each and suggest lines of research to obtain the evidence we lack. We then draw attention to a second major question, namely how changes in diet breadth affect the rest of a species’ biology. In particular, we know little about how changes in diet breadth feed back on genetic architecture, the population genetic environment, and other aspects of a species’ ecology. Knowing more about how generalists and specialists differ should go a long way toward sorting out potential explanations of specificity, and yield a deeper understanding of herbivorous insect diversity. Expected final online publication date for the Annual Review of Ecology, Evolution, and Systematics, Volume 51 is November 2, 2020. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
When herbivorous insects interact, they can increase or decrease each other's fitness. As it stands, we know little of what causes this variation. Classic competition theory predicts that competition will increase with niche overlap and population density. And classic hypotheses of herbivorous insect diversification predict that diet specialists will be superior competitors to generalists. Here, we test these predictions using phylogenetic meta‐analysis. We estimate the effects of diet breadth, population density and proxies of niche overlap: phylogenetic relatedness, physical proximity and feeding‐guild membership. As predicted, we find that competition between herbivorous insects increases with population density as well as phylogenetic and physical proximity. Contrary to predictions, competition tends to be stronger between than within feeding guilds and affects specialists as much as generalists. This is the first statistical evidence that niche overlap increases competition between herbivorous insects. However, niche overlap is not everything; complex feeding guild effects indicate important indirect interactions.
Pesticides and parasites have each been linked to increased mortality in western honey bees (Apis mellifera). Currently, it is uncertain if one makes the other worse; several studies have tested for potential synergistic stressor effects, but results have been mixed. Here, we use a hierarchical meta‐analysis of 63 experiments from 26 studies to gain a clearer view of the combined effects of parasites and pesticides on honey bee health. We found that combined pesticide–parasite treatments do tend to be deadlier than uncombined treatments but are significantly less deadly than predicted additive or multiplicative effects. In other words, combined treatment effects are not synergistic, but antagonistic. Much of the previous uncertainty about the combined effects of pesticides and parasites on honey bee health can be attributed to a bias in the previous research against stressor antagonism; many researchers have excluded the possibility of antagonism a priori. Synthesis and applications. Meta‐analysis shows that when honey bees are stressed by a combination of pesticides and parasites, the combined stress effect is antagonistic, that is, less than the sum of its parts. A better understanding of the mechanisms underlying this antagonism could prove critical for effective management of honey bee health.
Across herbivorous insect clades, species richness and host-use diversity tend to positively covary. This could be because host-use divergence drives speciation, or because it raises the ecological limits on species richness. To evaluate these hypotheses, we performed phylogenetic path model analyses of the species diversity of Nearctic aphids. Here, we show that variation in the species richness of aphid clades is caused mainly by host-use divergence, whereas variation in speciation rates is caused more by divergence in non-host-related niche variables. Aphid speciation is affected by both the evolution of host and non-host-related niche components, but the former is largely caused by the latter. Thus, our analyses suggest that host-use divergence can both raise the ecological limits on species richness and drive speciation, although in the latter case, host-use divergence tends to be a step along the causal path leading from non-host-related niche evolution to speciation.
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