Estimating the relative roles of top-down and bottom-up forces on insect herbivore populations: A classic study revisited

Although most ecologists agree that both top-down and bottom-up forces (predation and resource limitation, respectively) act in concert to influence populations of herbivores, it has proven difficult to estimate the relative contributions of such forces in terrestrial systems. Using a combination of time–series analysis of population counts recorded over 16 years and experimental data, we present the first estimates of the relative roles of top-down and bottom-up forces on the population dynamics of two terrestrial insect herbivores on the English oak (Quercus robur). Data suggest that temporal variation in winter moth, Operophtera brumata, density is dominated by time-lagged effects of pupal predators. By comparison, spatial variation in O. brumata density is dominated by host–plant quality. Overall, top-down forces explain 34.2% of population variance, bottom-up forces explain 17.2% of population variance, and 48.6% remains unexplained. In contrast, populations of the green oak tortrix, Tortrix viridana, appear dominated by bottom-up forces. Resource limitation, expressed as intraspecific competition among larvae for oak leaves, explains 29.4% of population variance. Host quality effects explain an additional 5.7% of population variance. We detected no major top-down effects on T. viridana populations. An unknown factor causing a linear decline in T. viridana populations over the 16-year study period accounts for most of the remaining unexplained variance. We discuss the observed differences between the insect species and the utility of time–series analysis as a tool in assessing the relative importance of top-down and bottom-up forces on herbivore populations.

Experimental evidence for a behavior-mediated trophic cascade in a terrestrial food chain

Predators of herbivorous animals can affect plant populations by altering herbivore density, behavior, or both. To test whether the indirect effect of predators on plants arises from density or behavioral responses in a herbivore population, we experimentally examined the dynamics of terrestrial food chains comprised of old field plants, leaf-chewing grasshoppers, and spider predators in Northeast Connecticut. To separate the effects of predators on herbivore density from the effects on herbivore behavior, we created two classes of spiders: (i) risk spiders that had their feeding mouth parts glued to render them incapable of killing prey and (ii) predator spiders that remained unmanipulated. We found that the effect of predators on plants resulted from predator-induced changes in herbivore behavior (shifts in activity time and diet selection) rather than from predator-induced changes in grasshopper density. Neither predator nor risk spiders had a significant effect on grasshopper density relative to a control. This demonstrates that the behavioral response of prey to predators can have a strong impact on the dynamics of terrestrial food chains. The results make a compelling case to examine behavioral as well as density effects in theoretical and empirical research on food chain dynamics.

Ape-like or hominid-like? The positional behavior of Oreopithecus bambolii reconsidered

Comparative morphological and functional analyses of the skeletal remains of Oreopithecus bambolii, a hominoid from the Miocene Mediterranean island of Tuscany–Sardinia (Italy), provides evidence that bipedal activities made up a significant part of the positional behavior of this primate. The mosaic pattern of its postcranial morphology is to some degree convergent with that of Australopithecus and functionally intermediate between apes and early hominids. Some unique traits could have been selected only under insular conditions where the absence of predators and the limitation of trophic resources play a crucial role in mammalian evolution.

Miriamin, a defensive diterpene from the eggs of a land slug (Arion sp.)

The eggs of the land slug Arion sp. contain a diterpene, miriamin, characterized as a polyoxygenated geranylgeraniol derivative. In bioassays with a coccinellid beetle, Harmonia axyridis, miriamin was shown to be potently antifeedant, indicating that the compound plays a protective role in nature. It is suggested that mucilaginous soil-inhabiting organisms, given their intense exposure to pathogens and predators, may be a rich source of chemical defensive agents.