Restricted dispersal reduces the strength of spatial density dependence in a tropical bird population

Spatial processes could play an important role in density-dependent population regulation because the disproportionate use of poor quality habitats as population size increases is widespread in animal populations-the so-called buffer effect. While the buffer effect patterns and their demographic consequences have been described in a number of wild populations, much less is known about how dispersal affects distribution patterns and ultimately density dependence. Here, we investigated the role of dispersal in spatial density dependence using an extraordinarily detailed dataset from a reintroduced Mauritius kestrel (Falco punctatus) population with a territorial (despotic) breeding system. We show that recruitment rates varied significantly between territories, and that territory occupancy was related to its recruitment rate, both of which are consistent with the buffer effect theory. However, we also show that restricted dispersal affects the patterns of territory occupancy with the territories close to release sites being occupied sooner and for longer as the population has grown than the territories further away. As a result of these dispersal patterns, the strength of spatial density dependence is significantly reduced. We conclude that restricted dispersal can modify spatial density dependence in the wild, which has implications for the way population dynamics are likely to be impacted by environmental change.

The population dynamical consequences of density-dependence in fungal plant pathogens

Almost all stages of a plant pathogen life cycle are potentially density dependent. At small scales and short time spans appropriate to a single-pathogen individual, density dependence can be extremely strong, mediated both by simple resource use, changes in the host due to defence reactions and signals between fungal individuals. In most cases, the consequences are a rise in reproductive rate as the pathogen becomes rarer, and consequently stabilisation of the population dynamics; however, at very low density reproduction may become inefficient, either because it is co-operative or because heterothallic fungi do not form sexual spores. The consequence will be historically determined distributions. On a medium scale, appropriate for example to several generations of a host plant, the factors already mentioned remain important but specialist natural enemies may also start to affect the dynamics detectably. This could in theory lead to complex (e.g. chaotic) dynamics, but in practice heterogeneity of habitat and host is likely to smooth the extreme relationships and make for more stable, though still very variable, dynamics. On longer temporal and longer spatial scales evolutionary responses by both host and pathogen are likely to become important, producing patterns which ultimately depend on the strength of interactions at smaller scales.

Declining survival rates in a reintroduced population of the Mauritius kestrel: evidence for non-linear density dependence and environmental stochasticity

1. We studied a reintroduced population of the formerly critically endangered Mauritius kestrel Falco punctatus Temmink from its inception in 1987 until 2002, by which time the population had attained carrying capacity for the study area. Post-1994 the population received minimal management other than the provision of nestboxes. 2. We analysed data collected on survival (1987-2002) using program MARK to explore the influence of density-dependent and independent processes on survival over the course of the population's development. 3.We found evidence for non-linear, threshold density dependence in juvenile survival rates. Juvenile survival was also strongly influenced by climate, with the temporal distribution of rainfall during the cyclone season being the most influential climatic variable. Adult survival remained constant throughout. 4. Our most parsimonious capture-mark-recapture statistical model, which was constrained by density and climate, explained 75.4% of the temporal variation exhibited in juvenile survival rates over the course of the population's development. 5. This study is an example of how data collected as part of a threatened species recovery programme can be used to explore the role and functional form of natural population regulatory processes. With the improvements in conservation management techniques and the resulting success stories, formerly threatened species offer unique opportunities to further our understanding of the fundamental principles of population ecology.

The effects of spatial and temporal heterogeneity on the population dynamics of four animal species in a Danish landscape

Background: Variation in carrying capacity and population return rates is generally ignored in traditional studies of population dynamics. Variation is hard to study in the field because of difficulties controlling the environment in order to obtain statistical replicates, and because of the scale and expense of experimenting on populations. There may also be ethical issues. To circumvent these problems we used detailed simulations of the simultaneous behaviours of interacting animals in an accurate facsimile of a real Danish landscape. The models incorporate as much as possible of the behaviour and ecology of skylarks Alauda arvensis, voles Microtus agrestis, a ground beetle Bembidion lampros and a linyphiid spider Erigone atra. This allows us to quantify and evaluate the importance of spatial and temporal heterogeneity on the population dynamics of the four species. Results: Both spatial and temporal heterogeneity affected the relationship between population growth rate and population density in all four species. Spatial heterogeneity accounted for 23–30% of the variance in population growth rate after accounting for the effects of density, reflecting big differences in local carrying capacity associated with the landscape features important to individual species. Temporal heterogeneity accounted for 3–13% of the variance in vole, skylark and spider, but 43% in beetles. The associated temporal variation in carrying capacity would be problematic in traditional analyses of density dependence. Return rates were less than one in all species and essentially invariant in skylarks, spiders and beetles. Return rates varied over the landscape in voles, being slower where there were larger fluctuations in local population sizes. Conclusion: Our analyses estimated the traditional parameters of carrying capacities and return rates, but these are now seen as varying continuously over the landscape depending on habitat quality and the mechanisms of density dependence. The importance of our results lies in our demonstration that the effects of spatial and temporal heterogeneity must be accounted for if we are to have accurate predictive models for use in management and conservation. This is an area which until now has lacked an adequate theoretical framework and methodology.

Effects of patch size and density on flower visitation and seed set of wild plants: a pan-European approach

1. Habitat fragmentation can affect pollinator and plant population structure in terms of species composition, abundance, area covered and density of flowering plants. This, in turn, may affect pollinator visitation frequency, pollen deposition, seed set and plant fitness. 2. A reduction in the quantity of flower visits can be coupled with a reduction in the quality of pollination service and hence the plants’ overall reproductive success and long-term survival. Understanding the relationship between plant population size and⁄ or isolation and pollination limitation is of fundamental importance for plant conservation. 3. Weexamined flower visitation and seed set of 10 different plant species fromfive European countries to investigate the general effects of plant populations size and density, both within (patch level) and between populations (population level), on seed set and pollination limitation. 4. Wefound evidence that the effects of area and density of flowering plant assemblages were generally more pronounced at the patch level than at the population level. We also found that patch and population level together influenced flower visitation and seed set, and the latter increased with increasing patch area and density, but this effect was only apparent in small populations. 5. Synthesis. By using an extensive pan-European data set on flower visitation and seed set we have identified a general pattern in the interplay between the attractiveness of flowering plant patches for pollinators and density dependence of flower visitation, and also a strong plant species-specific response to habitat fragmentation effects. This can guide efforts to conserve plant–pollinator interactions, ecosystem functioning and plant fitness in fragmented habitats.

Aggregation of parasitism risk in an aphid-parasitoid system: Effects of plant patch size and aphid density

Few studies have linked density dependence of parasitism and the tritrophic environment within which a parasitoid forages. In the non-crop plant-aphid, Centaurea nigra-Uroleucon jaceae system, mixed patterns of density-dependent parasitism by the parasitoids Aphidius funebris and Trioxys centaureae were observed in a survey of a natural population. Breakdown of density-dependent parasitism revealed that density dependence was inverse in smaller colonies but direct in large colonies (>20 aphids), suggesting there is a threshold effect in parasitoid response to aphid density. The CV2 of searching parasitoids was estimated from parasitism data using a hierarchical generalized linear model, and CV2>1 for A. funebris between plant patches, while for T. centaureae CV2>1 within plant patches. In both cases, density independent heterogeneity was more important than density-dependent heterogeneity in parasitism. Parasitism by T. centaureae increased with increasing plant patch size. Manipulation of aphid colony size and plant patch size revealed that parasitism by A. funebris was directly density dependent at the range of colony sizes tested (50-200 initial aphids), and had a strong positive relationship with plant patch size. The effects of plant patch size detected for both species indicate that the tritrophic environment provides a source of host density independent heterogeneity in parasitism, and can modify density-dependent responses. (c) 2007 Gessellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

The joint effects of larval density and C-14-cypermethrin on the life history and population growth rate of the midge Chironomus riparius

1. Chemical effects on organisms are typically assessed using individual-level endpoints or sometimes population growth rate (PGR), but such measurements are generally made at low population densities. In contrast most natural populations are subject to density dependence and fluctuate around the environmental carrying capacity as a result of individual competition for resources. As ecotoxicology aims to make reliable population projections of chemical impacts in the field, an understanding of how high-density or resource-limited populations respond to environmental chemicals is essential. 2. Our objective was to determine the joint effects of population density and chemical stress on the life history and PGR of an important ecotoxicological indicator species, Chironomus riparius, under controlled laboratory conditions. Populations were fed the same ration but initiated at different densities and exposed to a solvent control and three concentrations of C-14-cypermethrin in a sediment-water test system for 67 days at 20 +/- 1 degreesC. 3. Density had a negative effect on all the measured life-history traits, and PGR declined with increasing density in the controls. Exposure to C-14-cypermethrin had a direct negative effect on juvenile survival, presumably within the first 24 h because the chemical rapidly dissipated from the water column. Reductions in the initial larval densities resulted in an increase in the available resources for the survivors. Subsequently, exposed populations emerged sooner and started producing offspring earlier than the controls. C-14-cypermethrin had no effect on estimated fecundity and adult body weight but interacted with density to reduce the time to first emergence and first reproduction. As a result, PGR increased with cypermethrin concentration when populations were initiated at high densities. 4. Synthesis and applications. The results showed that the effects of C-14-cypermethrin were buffered at high density, so that the joint effects of density and chemical stress on PGR were less than additive. Low levels of chemical stressors may increase carrying capacity by reducing juvenile competition for resources. More and perhaps fitter adults may be produced, similar to the effects of predators and culling; however, toxicant exposure may result in survivors that are less tolerant to changing conditions. If less than additive effects are typical in the field, standard regulatory tests carried out at low density may overestimate the effects of environmental chemicals. Further studies over a wide range of chemical stressors and organisms with contrasting life histories are needed to make general recommendations.

The influence of larval density, food availability and habitat longevity on the life history and population growth rate of the midge Chironomus riparius

Despite long-standing interest in the forms and mechanisms of density dependence, these are still imperfectly understood. However, in a constant environment an increase in density must reduce per capita resource availability, which in turn leads to reduced survival, fecundity and somatic growth rate. Here we report two population experiments examining the density dependent responses under controlled conditions of an important indicator species, Chironomus riparius. The first experiment was run for 35 weeks and was started at low density with replicate populations being fed three different rations. Increased ration reduced generation time and increased population growth rate (pgr) but had no effect on survival, fecundity and female body weight in the first generation. In the second generation there was a six-fold increase in generation time, presumably due to the greatly reduced per capita resource availability as the estimated initial densities of the second generation were 300 times greater than the first. Juvenile survival to emergence, fecundity, adult body weight and pgr declined by 90%, 75%, 35% and 99%, respectively. These large between-generation effects may have obscured the effects of the threefold variation in ration, as only survival to emergence significantly increased with ration in the second generation. These results suggest that some chironomid larvae survive a reduction in resource availability by growing more slowly. In the ephemeral habitats sometimes occupied by C. riparius, the effects of population density may depend crucially on the longevity of the environment. A second experiment was therefore performed to measure pgr from six different starting densities over an eight-week period. The relationship between pgr and density was concave, viewed from above. At densities above 16 larvae per cm(2), less than 1% of the population emerged and no offspring were produced. Under the conditions of experiment 2 - an 8-week habitat lifespan carrying capacity was estimated as 8 larvae per cm(2).

The population dynamics of disease on short and long time-scales

Observational evidence is scarce concerning the distribution of plant pathogen population sizes or densities as a function of time-scale or spatial scale. For wild pathosystems we can only get indirect evidence from evolutionary patterns and the consequences of biological invasions.We have little or no evidence bearing on extermination of hosts by pathogens, or successful escape of a host from a pathogen. Evidence over the last couple of centuries from crops suggest that the abundance of particular pathogens in the spectrum affecting a given host can vary hugely on decadal timescales. However, this may be an artefact of domestication and intensive cultivation. Host-pathogen dynamics can be formulated mathematically fairly easily–for example as SIR-type differential equation or difference equation models, and this has been the (successful) focus of recent work in crops. “Long-term” is then discussed in terms of the time taken to relax from a perturbation to the asymptotic state. However, both host and pathogen dynamics are driven by environmental factors as well as their mutual interactions, and both host and pathogen co-evolve, and evolve in response to external factors. We have virtually no information about the importance and natural role of higher trophic levels (hyperpathogens) and competitors, but they could also induce long-scale fluctuations in the abundance of pathogens on particular hosts. In wild pathosystems the host distribution cannot be modelled as either a uniform density or even a uniform distribution of fields (which could then be treated as individuals). Patterns of short term density-dependence and the detail of host distribution are therefore critical to long-term dynamics. Host density distributions are not usually scale-free, but are rarely uniform or clearly structured on a single scale. In a (multiply structured) metapopulation with coevolution and external disturbances it could well be the case that the time required to attain equilibrium (if it exists) based on conditions stable over a specified time-scale is longer than that time-scale. Alternatively, local equilibria may be reached fairly rapidly following perturbations but the meta-population equilibrium be attained very slowly. In either case, meta-stability on various time-scales is a more relevant than equilibrium concepts in explaining observed patterns.

Graeme Caughley and the fundamentals of population ecology: a personal view

Caughley's contributions to single-species population ecology are here discussed in the light of some of the ideas and studies his work elicited, with particular reference to influences on my own work and that of my collaborators. Major themes are the manner and extent of population regulation; the alternate perspectives on population regulation that are obtained by density-dependence analyses and mechanistic analyses in terms of food availability and other causal factors; and ways in which mechanistic analyses can be elaborated to characterise a species' ecological niche and relate it to the species' geographic range.

Towards a population ecology of stressed environments: the effects of zinc on the springtail Folsomia candida

1. To understand population dynamics in stressed environments it is necessary to join together two classical lines of research. Population responses to environmental stress have been studied at low density in life table response experiments. These show how the population's growth rate (pgr) at low density varies in relation to levels of stress. Population responses to density, on the other hand, are based on examination of the relationship between pgr and population density. 2. The joint effects of stress and density on pgr can be pictured as a contour map in which pgr varies with stress and density in the same way that the height of land above sea level varies with latitude and longitude. Here a microcosm experiment is reported that compared the joint effects of zinc and population density on the pgr of the springtail Folsomia candida (Collembola). 3. Our experiments allowed the plotting of a complete map of the effects of density and a stressor on pgr. Particularly important was the position of the pgr= 0 contour, which suggested that carrying capacity varied little with zinc concentration until toxic levels were reached. 4. This prediction accords well with observations of population abundance in the field. The method also allowed us to demonstrate, simultaneously, hormesis, toxicity, an Allee effect and density dependence. 5. The mechanisms responsible for these phenomena are discussed. As zinc is an essential trace element the initial increase in pgr is probably a consequence of dietary zinc deficiency. The Allee effect may be attributed to productivity of the environment increasing with density at low density. Density dependence is a result of food limitation. 6. Synthesis and applications. We illustrate a novel solution based on mapping a population's growth rate in relation to stress and population density. Our method allows us to demonstrate, simultaneously, hormesis, toxicity, an Allee effect and density dependence in an important ecological indicator species. We hope that the approach followed here will prove to have general applicability enabling predictions of field abundance to be made from estimates of the joint effects of the stressors and density on population growth rate.

Non-testability of equal weights spatial dependence

We show that any invariant test for spatial autocorrelation in a spatial error or spatial lag model with equal weights matrix has power equal to size. This result holds under the assumption of an elliptical distribution. Under Gaussianity, we also show that any test whose power is larger than its size for at least one point in the parameter space must be biased.

Integrating spatial dependence into stochastic frontier analysis

An approach to incorporate spatial dependence into stochastic frontier analysis is developed and applied to a sample of 215 dairy farms in England and Wales. A number of alternative specifications for the spatial weight matrix are used to analyse the effect of these on the estimation of spatial dependence. Estimation is conducted using a Bayesian approach and results indicate that spatial dependence is present when explaining technical inefficiency.

How large is congressional dependence in agriculture. Bayesian inference about ‘scale’ and ‘scope’ in measuring a spatial externality

The political economy literature on agriculture emphasizes influence over political outcomes via lobbying conduits in general, political action committee contributions in particular and the pervasive view that political preferences with respect to agricultural issues are inherently geographic. In this context, ‘interdependence’ in Congressional vote behaviour manifests itself in two dimensions. One dimension is the intensity by which neighboring vote propensities influence one another and the second is the geographic extent of voter influence. We estimate these facets of dependence using data on a Congressional vote on the 2001 Farm Bill using routine Markov chain Monte Carlo procedures and Bayesian model averaging, in particular. In so doing, we develop a novel procedure to examine both the reliability and the consequences of different model representations for measuring both the ‘scale’ and the ‘scope’ of spatial (geographic) co-relations in voting behaviour.

Spatial variability of flow statistics within regular building arrays

Turbulence statistics obtained by direct numerical simulations are analysed to investigate spatial heterogeneity within regular arrays of building-like cubical obstacles. Two different array layouts are studied, staggered and square, both at a packing density of λp=0.25 . The flow statistics analysed are mean streamwise velocity ( u− ), shear stress ( u′w′−−−− ), turbulent kinetic energy (k) and dispersive stress fraction ( u˜w˜ ). The spatial flow patterns and spatial distribution of these statistics in the two arrays are found to be very different. Local regions of high spatial variability are identified. The overall spatial variances of the statistics are shown to be generally very significant in comparison with their spatial averages within the arrays. Above the arrays the spatial variances as well as dispersive stresses decay rapidly to zero. The heterogeneity is explored further by separately considering six different flow regimes identified within the arrays, described here as: channelling region, constricted region, intersection region, building wake region, canyon region and front-recirculation region. It is found that the flow in the first three regions is relatively homogeneous, but that spatial variances in the latter three regions are large, especially in the building wake and canyon regions. The implication is that, in general, the flow immediately behind (and, to a lesser extent, in front of) a building is much more heterogeneous than elsewhere, even in the relatively dense arrays considered here. Most of the dispersive stress is concentrated in these regions. Considering the experimental difficulties of obtaining enough point measurements to form a representative spatial average, the error incurred by degrading the sampling resolution is investigated. It is found that a good estimate for both area and line averages can be obtained using a relatively small number of strategically located sampling points.