William L. Vickery

Producers, scroungers, and group foraging

We have developed a model that reconciles information-sharing and producer-scrounger models of group foraging. Our model includes producers, scroungers, and an opportunistic forager that can both produce and scrounge but with reduced efficiency. We show that these three strategies can coexist only in the unlikely case that the opportunists loss in searching ability is exactly equal to its gain in scrounging ability. However, all pairs of strategies can coexist. Three parameters control the proportions of coexisting strategists: the degree of compatibility between the opportunists producing and scrounging activities; the proportion of food patches that are shared with scrounging individuals; and the effective group size. When there is little incompatibility between producing and scrounging, opportunists will always be present, unless the producer is able to consume most of the patch without sharing. The opportunist strategy is always excluded when there is a high degree of incompatibility between producing and scrounging. We consider the organismal and ecological factors that are likely to affect all three parameters. Our model predicts that scrounging behavior is likely to be selected in a wide range of foraging groups and that it may impose a considerable cost on sociality.

LANDSCAPE-SCALE DISTURBANCES AND CHANGES IN BIRD COMMUNITIES OF BOREAL MIXED-WOOD FORESTS

Bird community response to both landscape-scale and local (forest types) changes in forest cover was studied in three boreal mixed-wood forest landscapes modified by different types of disturbances: (1) a pre-industrial landscape where human settlement, agriculture, and logging activities date back to the early 1930s, (2) an industrial timber managed forest, and (3) a forest dominated by natural disturbances. Birds were sampled at 459 sampling stations distributed among the three landscapes. Local habitat and landscape characteristics of the context surrounding each sampling station (500-m and 1-km radius) were also computed. Bird communities were influenced by landscape-scale changes in forest cover. The higher proportion of early-successional habitats in both human-disturbed landscapes resulted in significantly higher abundance of early-successional bird species and generalists. The mean number of mature forest bird species was significantly lower in the industrial and pre-industrial landscapes than in the natural landscape. Landscape-scale conversion of mature forests from mixed-wood to deciduous cover in human-disturbed landscapes was the main cause of changes in mature forest bird communities. In these landscapes, the abundance of species associated with mixed and coniferous forest cover was lower, whereas species that preferred a deciduous cover were more abundant. Variation in bird community composition determined by the landscape context was as important as local habitat conditions, suggesting that predictions on the regional impact of forest management on songbirds with models solely based on local scale factors could be misleading. Patterns of bird species composition were related to several landscape composition variables (proportions of forest types), but not to configuration variables (e.g., interior habitat, amount of edge). Overall, our results indicated that the large-scale conversion of the southern portion of the boreal forest from a mixed to a deciduous cover may be one of the most important threats to the integrity of bird communities in these forest mosaics. Negative effects of changes in bird communities could be attenuated if current forestry practices are modified toward maintaining forest types (deciduous, mixed-wood, and coniferous) at levels similar to those observed under natural disturbances.

Detection of Density‐Dependent Effects in Annual Duck Censuses

Annual censuses of duck populations from two localities differing in environmental variability were used to test the effectiveness of six tests of density dependence. The magnitude of Type I error for each test was estimated from simulated density—independent data having the same mean and variance as the observed censuses. The actual census data were used to evaluate the ability to detect density dependence. Only autoregression had the desired level of type I error, but this test was ineffective when populations grew or declined. The standard major axis was effective in the latter cases but was ineffective when populations did not grow or decline. The only effective test applicable to all cases was a regression test with a rejection region based oin simulated data. Results of these latter analyses agree with earlier descriptions of interspecific variation in life—history tactics among duck species and the importance of competition as a factor affecting guild structure in prairie—nesting ducks. Diving ducks, which occupy the most temporally stable wetlands, show a greater degree of density dependence than do dabbling ducks, and competition influences the guild structure of diving ducks more than that of dabbling ducks.

Intratree Variation in Fruit Production and Implications for Primate Foraging

We tested the hypothesis that fruit quantity and quality vary vertically within trees. We quantified intratree fruit production before exploitation by frugivores at different heights in 89 trees from 17 species fed on by primates in Kibale National Park, Uganda. We also conducted a pilot study to determine if the nutritional value of fruit varied within tree crowns. Depending on the species and crown size, we divided tree canopies into 2 or 3 vertical layers. In 2-layered trees, upper crowns produced fruits that were 9.6–30.1% bigger and 0.52–140 times the densities of those from lower crowns, with one exception. Among 2-layered trees, upper crowns produced a mean of 46.9 fruits/m3 (median 12.1), while lower crowns produced a mean of 14.1 fruits/m3 (median 2.5). Among 3-layered trees, upper crowns produced a mean density of 49.9 fruits/m3 (median 12.5), middle crowns a mean of 16.8 fruits/m3 (median 6.6), and lower crowns a mean of 12.8 fruits/m3 (median 1.8). Dry pulp and moisture were systematically greater per fruit in the highest compared to the lowest canopy layers (22.4% and 16.4% respectively in 2-layered trees, 49.7% and 21.8% respectively in 3-layered trees). In 1 tree of Diospyros abyssinica, a pilot nutritional study showed that upper crown ripe fruit contained 41.9% more sugar, 8.4% more crude proteins, and 1.8 times less of the potentially toxic saponin than lower crown ripe fruit, but the result needs to be verified with more individuals and species of trees. We discuss the consequences of intratree variations in fruit production with respect to competition among frugivorous primates.

Parasite-mediated sexual selection: just how choosy are parasitized females?

Abstract Models of parasite-mediated sexual selection have thus far overlooked the potential effects of parasites of females on their hosts’ ability to choose mates. A set of models addressing this issue is developed, each building on the previous one to add complexity and realism to the framework. The selection coefficient for parasite immunity and brightness is estimated using the ratio of the fitness of susceptible males to the fitness of immune males. Parasite-induced reduction in female choosiness can substantially relax the selection for bright, immune males, especially when: (1) immunity to parasites is rare in the population, (2) parasites are not highly aggregated within the host population, (3) parasites are abundant, and (4) the effects of parasites on male brightness or female choosiness are severe. Parasite-induced variability in male brightness is most likely to occur in populations in which parasites are abundant and not aggregated; if females in those populations show a reduced preference for bright males, sexual selection for brightness (and parasite immunity) will still operate but exert a weaker selective pressure.

Tree Climbing Strategies for Primate Ecological Studies

Primate ecological studies can benefit from accessing the canopy to estimate intra-tree and inter-tree variation in food availability and nutrient value, patch and subpatch depletion, foraging efficiency, as well as nest structure and nesting behaviors, parasitic transmission and predator detectability. We compare several ways to access the canopy and examine their suitability for studies of primates. Two of them—the Single Rope Technique and the Climbing Spur Method—allow people to safely access almost all kinds of trees, regardless of their size, height or shape. Modern climbing gear and contemporaneous safety protocols, derived from rock climbers, speleologists, and industrial arborists, are reliable and appropriate for primate ecological studies. Climbing gear is specialized and still expensive for students, but tree climbing can be dangerous during specific maneuvres. Consequently, formal training and preliminary experience are essential before attempting to collect data. We discuss the physics of falling, risk assessment associated with a fall, knots, gear and safety precautions. Finally, we propose a Tree Climbing Safety Protocol adapted for 2 climbing methods and primate field ecology. Researchers should be aware that climbing safety depends on their own judgment, which must be based on competent instruction, experience, and a realistic assessment of climbing ability. Therefore, the information we provide should be used only to supplement competent personal instruction and training in situ. Although most primate observations have been and will mostly be done from the ground in the future, canopy information complements the observations. Canopy data will add a significant new dimension to our knowledge of primates by providing strategic information otherwise unavailable.

Testing for density-dependent effects in sequential censuses

SummaryIn response to Gaston and Lawton (1987), we evaluated the ability of four statistical procedures to detect density dependence. We used data from the same 16 populations as Gaston and Lawton (1987). In each population, density dependence had been previously established with techniques that use more extensive data. The major axis test (Slade 1977) was rarely (3 populations of 16) capable of detecting density dependence. The autocorrelation test (Bulmer 1975) detected density dependence in 5 of 16 species (14 of 59 tests overall). The randomization procedure (Pollard et al. 1987) detected density dependence in 7 of the 16 data sets (10 of 59 tests overall). The simulation procedure (Vickery and Nudds 1984) detected density dependence in 5 of the 16 data sets (11 of 59 tests overall). We suggest that not all annual census data taken from populations subject to density-dependent effects will actually show evidence of such effects. We conclude that Pollard et al. s (1987) randomization procedure is the best test for detecting density dependence in sequential census data but it is not as powerful as more elaborate techniques (k-factor analysis, experimentation, etc.), nor is it meant to replace more extensive analyses.

Parasite extinction and colonisation and the evolution of parasite communities: a simulation study.

We determined what evolutionary processes influence the likelihood of detecting an effect of host ecological characteristics on the richness of parasite communities in comparative analyses among related host species. We used a mathematical model to generate phylogenies of hosts in which parasite communities varied over evolutionary time as parasite species were either gained or lost during host speciation events. Gain or loss of parasites were stochastic and could either be strongly, moderately, weakly, or not, affected by host ecological characteristics. The model was evaluated over this range of effects of host ecology, and for various mean probabilities of parasite gain and loss and various rates of change in host ecological characteristics at speciation events. Our results suggest that phylogenetic effects (the passing of parasite species from mother to daughter host species) are likely to obscure ecological effects (the effect of host body size, diet, habitat, lifespan, etc.) except when the effects of host ecology are strong, and the probabilities of gain or loss of parasites are high, or host ecological characteristics change markedly at speciation events. This outcome was not influenced by the shape of the phylogenetic tree used in the simulations. Sensitivity analysis of our model also shows this result to be robust to a wide range of assumptions and parameter values. Thus, because the composition of parasite communities tends to reflect their ancestry, the effect of host ecology will often be very difficult to detect.

TERRITORIALITY, VEGETATION COMPLEXITY, AND BIOLOGICAL CONTROL: THE CASE FOR SPIDERS

We constructed a model of territoriality relating the probability of starvation in a nonstationary, variable environment to the costs and benefits of feeding, defense, and exploration and their variances. The model suggests that survival in a territory increases with increasing food abundance and with decreasing defense and exploration costs. Territoriality, a risk-averse behavior, is favored when an animals net expected energy balance is positive (see also Stephens 1981). When the expected energy balance is negative, risk-prone behavior, abandoning the territory to become a marginal or floater, is appropriate. This result generalizes that of Stephens (1981) to the more general circumstances of nonstationary random environments. Our model allows discussion of the impact of habitat diversity on the density of territorial animals and on the control of pest populations. By stabilizing the variance of herbivore abundance, greater crop diversity should increase both survival and packing of the herbivores territorial predators. This should maximize herbivore consumption by territorial predators (spiders) and thus increase their contribution to the control of the phytophage population in agroecosystems.

The evolution of host manipulation by parasites: a game theory analysis

Many parasites are known to manipulate the behaviour of intermediate hosts in order to increase their probability of transmission to definitive hosts. This manipulation must have costs. Here we explore the combined effects of three such costs on the amount of effort a parasite should expend on host manipulation. Manipulation can have direct costs to future reproductive success due to energy expended to manipulate the host. There may also be indirect costs if other parasites infect the host and profit from the manipulation without paying the cost of manipulation. These “free riders” may impose a third cost by competing with manipulators for resources within the host. Using game theory analysis and several different competition models we show that intrahost competition will decrease the investment that a parasite should make in manipulation but that manipulation can, under some circumstances, be a profitable strategy even in the presence of non-manipulating competitors. The key determinants of the manipulator’s success and its investment in manipulation are the relatedness among parasites within the host, the ratio of the passive transmission rate to the efficiency of increasing transmission rate and the strength of competitive effects. Manipulation, when exploited by others, becomes an altruistic behaviour. Thus we suggest that our model may be generally applicable to cases where organisms can exploit the investment of others (possibly kin) while also competing with the organism whose investment they exploit.