Douglas J. Levey

The life histories of many species depend first on dispersal to local sites and then on establishment. After dispersal, density‐independent and density‐dependent mortalities modify propagule supply, determining the number of individuals that establish. Because multiple factors influence recruitment, the dichotomy of propagule versus establishment limitation is best viewed as a continuum along which the strength of propagule or establishment limitation changes with propagule input. To evaluate the relative importance of seed and establishment limitation for plants, we (1) describe the shape of the recruitment function and (2) use limitation and elasticity analyses to quantify the sensitivity of recruitment to perturbations in seed limitation and density‐independent and density‐dependent mortality. Using 36 seed augmentation studies for 18 species, we tested four recruitment functions against one another. Although the linear model (accounting for seed limitation and density‐independent mortality) fitted the largest number of studies, the nonlinear Beverton‐Holt model (accounting for density‐dependent mortality) performed better at high densities of seed augmentation. For the 18 species, seed limitation constrained population size more than other sources of limitation at ambient conditions. Seedling density reached saturation with increasing seed density in many studies, but at such high densities that seedling density was primarily limited by seed availability rather than microsite availability or density dependence.

The directed deterrence hypothesis posits that secondary metabolites in ripe fruit function to deter fruit consumption by vertebrates that do not disperse seeds, while not impacting consumption by those that do. We tested this hypothesis in two species of wild chilies (Capsicum spp.). Both produce fruits that contain capsaicinoids, the compounds responsible for the pungency of chilies. Previous work suggests seed-dispersing birds but not seed-destroying rodents consume chili fruits, presumably because rodents are deterred by capsaicin. However, fruit removal from chili plants by rodents and other mammals has not been previously explored. Because laboratory rodents can develop a preference for capsaicin, it is quite possible that wild rodents are natural consumers of chili fruits. We monitored the fate of 125 marked fruits of Capsicum chacoense and 291 fruits of Capsicum annuum. For both species, essentially all fruit removal occurred during the day, when rodents are inactive. Video monitoring revealed fruit removal only by birds, mostly by species known to disperse chili seeds in viable condition. Furthermore, these species are from taxonomic groups that tend to specialize on lipid-rich fruits. Both species of chili produce fruits that are unusually high in lipids (35% in C. chacoense, 24% in C. annuum). These results support the directed deterrence hypothesis and suggest that fruiting plants distinguish between seed predators and seed dispersers by producing fruits that repel the former and attract the latter.

Habitat fragmentation threatens biodiversity by disrupting dispersal. The mechanisms and consequences of this disruption are controversial, primarily because most organisms are difficult to track. We examined the effect of habitat corridors on long-distance dispersal of seeds by birds, and tested whether small-scale (<20 meters) movements of birds could be scaled up to predict dispersal of seeds across hundreds of meters in eight experimentally fragmented landscapes. A simulation model accurately predicted the observed pattern of seed rain and revealed that corridors functioned through edge-following behavior of birds. Our study shows how models based on easily observed behaviors can be scaled up to predict landscape-level processes.

Summary: Frugivorous vertebrates may affect plant fitness by dispersing seeds to sites favourable for establishment and by passing seeds through their gut, thereby altering germination patterns. Although gut passage can inhibit germination, most studies have found that it improves germination rate and/or success. However, studies that compare seeds passed through a gut to seeds directly removed from a fruit cannot quantify the overall effect of gut passage on seed germination. They address the ecologically narrow question of whether mechanical and/or chemical action in the gut affects germination. They do not address the germination potential of seeds in unconsumed fruits and therefore ignore a common seed fate, deposition in a fruit. We surveyed 99 studies that included the effect of vertebrate gut passage on germination and found that only 22 included germination from intact fruits. We suggest that the strongest experimental design to evaluate the impact of gut passage should include intact fruits, since release from germination inhibitors and high osmotic pressure are mechanisms that can alter germination.

Corridors connect otherwise isolated habitat patches and can direct movement of animals among such patches. In eight experimental landscapes, we tested two hypotheses of how corridors might affect dispersal behavior. The Traditional Corridor hypothesis posits that animals preferentially leave patches via corridors, following them into adjacent patches. The Drift Fence hypothesis posits that animals dispersing through matrix habitat are diverted into patches with corridors because they follow corridors when encountered. House flies (Musca domestica L.), a species that prefers the habitat of our patches and corridors, were released in a central patch (100×100 m) and recaptured in peripheral patches that were or were not connected by a corridor. Flies were captured more frequently in connected than unconnected patches, thereby supporting the Traditional Corridor hypothesis. The Drift Fence hypothesis was also supported, as flies were captured more frequently in unconnected patches with blind (dead end) corridors than in unconnected patches of equal area without blind corridors. A second experiment tested whether these results might be dependent on the type of patch-matrix boundary encountered by dispersing flies and whether edge-following behavior might be the mechanism underlying the observed corridor effect in the first experiment. We recorded dispersal patterns of flies released along forest edges with dense undergrowth in the forest (“closed” edges) and along edges with little forest understory (“open” edges). Flies were less likely to cross and more likely to follow closed edges than open edges, indicating that when patch and corridor edges are pronounced, edge-following behavior of flies may direct them along corridors into connected patches. Because edges in the first experiment were open, these results also suggest that corridor effects for flies in that experiment would have been even stronger if the edges around the source patches and corridors had been more closed. Taken together, our results suggest that corridors can affect dispersal of organisms in unappreciated ways (i.e., as drift fences) and that edge type can alter dispersal behavior.

Reports of tool usage by birds tend to be anecdotal as only a few individuals may be involved and the behaviour observed can often be interpreted in other ways. Here we describe the widespread collection of mammalian dung by burrowing owls (Athene cunicularia) and show that they use this dung as a bait to attract dung beetles, a major item of prey. Our controlled investigation provides an unambiguous estimate of the importance of tool use in a wild animal.

Understanding the evolutionary ecology of ethanol production may yield insights into why humans are prone to excessive consumption of ethanol. In particular, Dudley (2000) suggested that human ancestors developed a genetically based attraction to ethanol because they could use its odor plume to locate fruiting trees and because of health benefits from its consumption. If so, ethanol should be common in wild fruits and frugivores should prefer fruits with higher ethanol content. A literature review reveals that ethanol is indeed common in wild fruits but that it typically occurs in very low concentrations. Furthermore, frugivores strongly prefer ripe over rotting fruits, even though the latter may contain more ethanol. (Data on ethanol content of ripe and rotting wild fruit are lacking.) These results cast doubt on Dudley's hypothesis and raise the question of how humans became exposed to sufficiently high concentrations of ethanol to allow its excessive consumption. Because fermentation is an ancient and widespread practice, I suggest that humans “discovered” ethanol while using fermentation as a food preservation technique. They may have been predisposed to consume ethanol from previous and beneficial exposure to much lower doses or they may have become addicted to it at high concentrations because of fortuitous physiological responses.

Modulation of gut function is important in an ecological and evolutionary context because it likely determines what food items an animal can and cannot eat. We examined how diet affects activity of digestive enzymes in an omnivorous bird, the pine warbler (Dendroica pinus). Pine warblers were fed insect-based, fruit-based, and seed-based diets for approximately 54 d. We then measured activity of amylase, maltase, sucrase, aminopeptidase-N, trypsin, chymotrypsin, carboxypeptidase A, carboxypeptidase B, pancreatic lipase, and carboxyl ester lipase. We predicted that carbohydrase activities would be highest in birds fed the diet highest in carbohydrates (fruit based), protease activities would be highest in those fed the diet highest in protein (insect based), and lipase activities would be highest in those fed the diets highest in lipid (insect based and seed based). Also, we predicted that pine warblers would exhibit greater dietary modulation of enzyme activity than reported for a less omnivorous congener, the yellow-rumped warbler (Dendroica coronata). All predictions were upheld, supporting the hypothesis that pine warblers modulate the activity of digestive enzymes in proportion to demand from substrates in the diet.