Consumptive and nonconsumptive effects of predators on metacommunities of competing prey

Although predators affect prey both via consumption and by changing prey migration behavior, the interplay between these two effects is rarely incorporated into spatial models of predator-prey dynamics and competition among prey. We develop a model where generalist predators have consumptive effects (i.e., altering the likelihood of local prey extinction) as well as nonconsumptive effects (altering the likelihood of colonization) on spatially separated prey populations (metapopulations). We then extend this model to explore the effects of predators on competition among prey. We find that generalist predators can promote persistence of prey metapopulations by promoting prey colonization, but predators can also hasten system-wide extinction by either increasing local extinction or reducing prey migration. By altering rates of prey migration, predators in one location can exert remote control over prey dynamics in another location via predator-mediated changes in prey flux. Thus, the effect of predators may extend well beyond the proportion of patches they visit. In the context of prey metacommunities, predator-mediated shifts in prey migration and mortality can shift the competition-colonization trade-off among competing prey, leading to changes in the prey community as well as changes in the susceptibility of prey species to habitat loss. Consequently, native prey communities may be susceptible to invasion not only by exotic prey species that experience reduced amounts of mortality from resident predators, but also by exotic prey species that exhibit strong dispersal in response to generalist native predators. Ultimately, our work suggests that the consumptive and nonconsumptive effects of generalist predators may have strong, yet potentially cryptic, effects on competing prey capable of mediating coexistence, fostering invasion, and interacting with anthropogenic habitat alteration.     

Emergence cues of a mayfly in a high-altitude stream ecosystem: potential response to climate change.

To understand the consequences of human accelerated environmental change, it is important to document the effects on natural populations of an increasing frequency of extreme climatic events. In stream ecosystems, recent climate change has resulted in extreme variation in both thermal and hydrological regimes. From 2001 to 2004, a severe drought in western United States corresponded with earlier emergence of the adult stage of the high-altitude stream mayfly, Baetis bicaudatus. Using a long-term database from a western Colorado stream, the peak emergence date of this mayfly population was predicted by both the magnitude and date of peak stream flow, and by the mean daily water temperature, suggesting that Baetis may respond to declining stream flow or increasing water temperature as proximate cues for early metamorphosis. However, in a one-year survey of multiple streams from the same drainage basin, only water temperature predicted spatial variation in the onset of emergence of this mayfly. To decouple the effects of temperature and flow, we separately manipulated these factors in flow-through microcosms and measured the timing of B. bicaudatus metamorphosis to the adult stage. Mayflies emerged sooner in a warmed-water treatment than an ambient-water treatment; but reducing flow did not accelerate the onset of mayfly emergence. Nonetheless, using warming temperatures to cue metamorphosis enables mayflies to time their emergence during the descending limb of the hydrograph when oviposition sites (protruding rocks) are becoming available. We speculate that large-scale climate changes involving warming and stream drying could cause significant shifts in the timing of mayfly metamorphosis, thereby having negative effects on populations that play an important role in stream ecosystems.     

From individuals to ecosystem function: toward an integration of evolutionary and ecosystem ecology

An important goal in ecology is developing general theory on how the species composition of ecosystems is related to ecosystem properties and functions. Progress on this front is limited partly because of the need to identify mechanisms controlling functions that are common to a wide range of ecosystem types. We propose that one general mechanism, rooted in the evolutionary ecology of all species, is adaptive foraging behavior in response to predation risk. To support our claim, we present two kinds of empirical evidence from plant-based and detritus-based food chains of terrestrial and aquatic ecosystems. The first kind comes from experiments that explicitly trace how adaptive foraging influences ecosystem properties and functions. The second kind comes from a synthesis of studies that individually examine complementary components of particular ecosystems that together provide an integrated perspective on the link between adaptive foraging and ecosystem function. We show that the indirect effects of predators on plant diversity, plant productivity, nutrient cycling, trophic transfer efficiencies, and energy flux caused by consumer foraging shifts in response to risk are qualitatively different from effects caused by reductions in prey density due to direct predation. We argue that a perspective of ecosystem function that considers effects of consumer behavior in response to predation risk will broaden our capacity to explain the range of outcomes and contingencies in trophic control of ecosystems. This perspective also provides an operational way to integrate evolutionary and ecosystem ecology, which is an important challenge in ecology.     

Framing the search for a theory of land use

ABSTRACT

Land system science and affiliated research linked to sustainability require improved understanding and theorization of land and its change as a social-ecological system (SES). The absence of a general land-use theory, anchored in the social subsystem but with explicit links to the environmental subsystem, hampers this effort. Drawing on land-use explanations, meta-analyses, and associated frameworks, we advance a broad framework structure of eight elements – aggregations of explanatory variables – with links to the biophysical subsystem, for systematic comparisons of extant explanations. Tests and models can be employed to identify which set of variables and their configurations provide robust explanations of across land uses, identifying the potential for theory development. The framework and its application are applicable to both top-down and bottom-up explanatory approaches employed in the social sciences. Links to the environmental subsystem invite future exploration of SES explanations that reach across the different dimensions of global change and sustainability science.     

Revisiting the classics: considering nonconsumptive effects in textbook examples of predator-prey interactions

Predator effects on prey dynamics are conventionally studied by measuring changes in prey abundance attributed to consumption by predators. We revisit four classic examples of predator-prey systems often cited in textbooks and incorporate subsequent studies of nonconsumptive effects of predators (NCE), defined as changes in prey traits (e.g., behavior, growth, development) measured on an ecological time scale. Our review revealed that NCE were integral to explaining lynx-hare population dynamics in boreal forests, cascading effects of top predators in Wisconsin lakes, and cascading effects of killer whales and sea otters on kelp forests in nearshore marine habitats. The relative roles of consumption and NCE of wolves on moose and consequent indirect effects on plant communities of Isle Royale depended on climate oscillations. Nonconsumptive effects have not been explicitly tested to explain the link between planktonic alewives and the size structure of the zooplankton, nor have they been invoked to attribute keystone predator status in intertidal communities or elsewhere. We argue that both consumption and intimidation contribute to the total effects of keystone predators, and that characteristics of keystone consumers may differ from those of predators having predominantly NCE. Nonconsumptive effects are often considered as an afterthought to explain observations inconsistent with consumption-based theory. Consequently, NCE with the same sign as consumptive effects may be overlooked, even though they can affect the magnitude, rate, or scale of a prey response to predation and can have important management or conservation implications. Nonconsumptive effects may underlie other classic paradigms in ecology, such as delayed density dependence and predator-mediated prey coexistence. Revisiting classic studies enriches our understanding of predator-prey dynamics and provides compelling rationale for ramping up efforts to consider how NCE affect traditional predator-prey models based on consumption, and to compare the relative magnitude of consumptive and NCE of predators.     

Swarming and mating behavior of a mayfly Baetis bicaudatus suggest stabilizing selection for male body size

Large size often confers a fitness advantage to female insects because fecundity increases with body size. However, the fitness benefits of large size for male insects are less clear. We investigated the mating behavior of the mayfly Baetis bicaudatus to determine whether the probability of male mating success increased with body size. Males formed mating aggregations (swarms) ranging from a few to hundreds of individuals, 1-4 m above the ground for about 1.5-2 h in the early morning. Females that flew near swarms were grabbed by males, pairs dropped to the vegetation where they mated and then flew off individually. Some marked males returned to swarms 1, 2 or 3 days after marking. Larger males swarmed near spruce trees at the edges of meadows, but the probability of copulating was not a function of male body size (no large male advantage). Furthermore, the potential fitness advantage of mating with larger, more fecund females was not greater for large males (no size-assortative mating). However, the sizes of copulating males were significantly less variable than those of non-mating males collected at random in swarms. Intermediate male size may be optimal during mating because of trade-offs between flight agility and longevity or competitive ability. Results of this study are consistent with the hypotheses that there is stabilizing selection on adult male body size during mating, and that male body size in this species may be influenced more by selection pressures acting on larvae than on adults.     

A comparative study of the costs of alternative mayfly oviposition behaviors

Oviposition behavior of insects has associated fitness costs related to the probability that females survive to oviposit. During summer 2003, we observed the oviposition behavior and compared the mortality rates of females of 17 mayfly species in one western Colorado watershed. We dissected adult females collected on terrestrial sticky traps, in drift nets submerged in streams, and in stomachs of brook trout to determine whether the mayflies had oviposited before capture, drowning, or consumption. Females oviposited by either splashing on the water surface releasing all their eggs (splashers), dropping their eggs from the air (bombers), dipping their abdomens multiple times releasing a few eggs at a time (dippers), landing on rocks and ovipositing on the undersides (landers), or floating downstream while releasing their eggs (floaters). Almost 100% of lander and 50% of dipper females had not oviposited when captured on sticky traps, increasing their vulnerability to preoviposition mortality by aerial predators compared to mayflies with other behaviors. In contrast, most females had laid their eggs before drowning or being eaten by a fish (50–90%). However, groups with oviposition behaviors most exposed to the water surface (floaters, then splashers, dippers, and landers) were more vulnerable to drowning before completing oviposition. In addition, splashers and floaters were most vulnerable to predation by brook trout before ovipositing. These data suggest that fitness costs associated with preoviposition mortality may be considerable depending on mayfly oviposition behavior. Furthermore, previously demonstrated benefits of low predation rates on eggs of lander species may be offset in part by costs to female survival. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.