Spatial scale of local breeding habitat quality and adjustment of breeding decisions

Experimental studies provide evidence that, in spatially and temporally heterogeneous environments, individuals track variation in breeding habitat quality to adjust breeding decisions to local conditions. However, most experiments consider environmental variation at one spatial scale only, while the ability to detect the influence of a factor depends on the scale of analysis. We show that different breeding decisions by adults are based on information about habitat quality at different spatial scales. We manipulated (increased or decreased) local breeding habitat quality through food availability and parasite prevalence at a small (territory) and a large (patch) scale simultaneously in a wild population of Great Tits (Parus major). Females laid earlier in high-quality large-scale patches, but laying date did not depend on small-scale territory quality. Conversely, offspring sex ratio was higher (i.e., biased toward males) in high-quality, small-scale territories but did not depend on large-scale patch quality. Clutch size and territory occupancy probability did not depend on our experimental manipulation of habitat quality, but territories located at the edge of patches were more likely to be occupied than central territories. These results suggest that integrating different decisions taken by breeders according to environmental variation at different spatial scales is required to understand patterns of breeding strategy adjustment.     

Local population size in a flightless insect: importance of patch structure-dependent mortality

In spatially heterogeneous systems, utilizing population models to integrate the effects of multiple population rates can yield powerful insights into the relative importance of the component rates. The relative importance of demographic rates and dispersal in shaping the distribution of the western tussock moth (Orgyia vetusta) among patches of its host plant was explored using stage-structured population models. Tussock moth dispersal occurs passively in first-instar larvae and is poor or absent in all other life stages. Spatial surveys suggested, however, that moth distribution is not well explained by passive dispersal; moth populations were greater on small patches and on isolated ones. Further analysis showed that several local demographic rates varied significantly with patch characteristics. Two mortality factors in particular may explain the observed patterns. First, crawler mortality both increased with patch size and was density-dependent. A single-patch difference equation model showed mortality related to patch size is strong enough to overcome the homogenizing effect of density dependence; greater equilibrium densities were predicted for smaller patches. Second, although three rates were found to vary with local patch density, only pupal parasitism by a chalcid wasp could potentially account for higher moth abundances on isolated patches. A spatially explicit simulation model of the multiple-patch system showed that spatial variation in pupal parasitism is indeed strong enough to generate such a pattern. These results demonstrate that habitat spatial structure can affect multiple population processes simultaneously, and even relatively low attack rates imposed on a reproductively valuable life stage of the host can have a dominant effect on population distribution among habitat patches.     

Can non-breeding be a cost of breeding dispersal?

Breeding habitat selection and dispersal are crucial processes that affect many components of fitness. Breeding dispersal entails costs, one of which has been neglected: dispersing animals may miss breeding opportunities because breeding dispersal requires finding a new nesting site and mate, two time- and energy-consuming activities. Dispersers are expected to be prone to non-breeding. We used the kittiwake (Rissa tridactyla) to test whether breeding dispersal influences breeding probability. Breeding probability was associated with dispersal, in that both were negatively influenced by private information (previous individual reproductive success) and public information (average reproductive success of conspecifics) about patch quality. Furthermore, the probability of skipping breeding was 1.7 times higher in birds that settled in a new patch relative to those that remained on the same patch. Finally, non-breeders that resumed breeding were 4.4 times more likely to disperse than birds that bred in successive years. Although private information may influence breeding probability directly, the link between breeding probability and public information may be indirect, through the influence of public information on breeding dispersal, non-breeding thus being a cost of dispersal. These results support the hypothesis that dispersal may result in not being able to breed. More generally, non-breeding (which can be interpreted as an extreme form of breeding failure) may reveal costs of various previous activities. Because monitoring the non-breeding portion of a population is difficult, non-breeders have been neglected in many studies of reproduction trade-offs.     

Do source-sink dynamics promote the spread of an invasive grass into a novel habitat?

Models of source-sink and other spatial patch dynamics have generated a number of ideas and predictions about species range expansion, the evolution of local adaptation, and the factors influencing population persistence, but relatively few empirical studies have applied these ideas due to the difficulty of measuring both patch-specific demography and movement rates. In this study, I used a combination of mark-recapture experiments, model fitting, and demographic approaches to ask how habitat-specific differences in population growth and dispersal affect spread of the invasive grass Aegilops triuncialis into serpentine environments. A. triuncialis germinated at lower rates but exhibited equivalent survival and greater growth in edge (extreme serpentine) than in core populations, even accounting for density differences between habitats. Estimated growth rates (lambda) for four of five edge subpopulations were strongly positive, ranging from lambda = 1.32 to 2.09 without propagule input from adjacent habitat. Local dispersal was best described by an exponential kernel, with a mean dispersal distance about twice as long on the edge (0.24-0.40 m) as in the core (0.18 m). Twenty-five percent of marked spikes in the edge were not relocated within the patch, suggesting greater rates of either seed predation or long-distance dispersal that reduced population growth. These results suggest that A. triuncialis can successfully spread into extreme serpentine habitats without sustained propagule input from adjacent populations. Further, asymmetric dispersal that may be both habitat- and density-dependent could slow growth rates on the edge. This pattern may also increase the importance of harsh edge patches as a source of long-distance dispersers.     

Dispersal depression with habitat fragmentation in the bog fritillary butterfly

Habitat fragmentation is expected to impose strong selective pressures on dispersal rates. However, evolutionary responses of dispersal are not self-evident, since various selection pressures act in opposite directions. Here we disentangled the components of dispersal behavior in a metapopulation context using the Virtual Migration model, and we linked their variation to habitat fragmentation in the specialist butterfly Proclossiana eunomia. Our study provided a nearly unique opportunity to study how habitat fragmentation modifies dispersal at the landscape scale, as opposed to microlandscapes or simulation studies. Indeed, we studied the same species in four landscapes with various habitat fragmentation levels, in which large amounts of field data were collected and analyzed using similar methodologies. We showed the existence of quantitative variations in dispersal behavior correlated with increased fragmentation. Dispersal propensity from habitat patches (for a given patch size), and mortality during dispersal (for a given patch connectivity) were lower in more fragmented landscapes. We suggest that these were the consequences of two different evolutionary responses of dispersal behavior at the individual level: (1) when fragmentation increased, the reluctance of individuals to cross habitat patch boundaries also increased; (2) when individuals dispersed, they flew straighter in the matrix, which is the best strategy to improve dispersal success. Such evolutionary responses could generate complex nonlinear patterns of dispersal changes at the metapopulation level according to habitat fragmentation. Due to the small size and increased isolation of habitat patches in fragmented landscapes, overall emigration rate and mortality during dispersal remained high. As a consequence, successful dispersal at the metapopulation scale remained limited. Therefore, to what extent the selection of individuals with a lower dispersal propensity and a higher survival during dispersal is able to limit detrimental effects of habitat fragmentation on dispersal success is unknown, and any conclusion that metapopulations would compensate for them is flawed.     

Modelling dispersal with diffusion and habitat selection: Analytical results for highly fragmented landscapes

Quantifying dispersal is crucial both for understanding ecological population dynamics, and for gaining insight into factors that affect the genetic structure of populations. The role of dispersal becomes pronounced in highly fragmented landscapes inhabited by spatially structured populations. We consider a landscape consisting of a set of habitat patches surrounded by unsuitable matrix, and model dispersal by assuming that the individuals follow a random walk with parameters that may be specific to the habitat type. We allow for spatial variation in patch quality, and account for edge-mediated behavior, the latter meaning that the individuals bias their movement towards the patches when close to an edge between a patch and the matrix. We employ a diffusion approximation of the random walk model to derive analytical expressions for various characteristics of the dispersal process. For example, we derive formulae for the time that an individual is expected to spend in its current patch i, and for the time that it will spend in the matrix, both conditional on the individual hitting next a given patch j before hitting any of the other patches or dying. The analytical formulae are based on the assumptions that the landscape is infinitely large, that the patches are circularly shaped, and that the patches are small compared to interpatch distances. We evaluate the effect of these assumptions by comparing the analytical results to numerical results in a real patch network that violates all of the three assumptions. We then consider a landscape that fulfills the assumptions, and show that in this case the analytical results are in a very good agreement with the numerical results. The results obtained here allow the construction of computationally efficient dispersal models that can be used as components of metapopulation models.     

Quantifying the importance of patch-specific changes in habitat to metapopulation viability of an endangered songbird

A growing number of programs seek to facilitate species conservation using incentive-based mechanisms. Recently, a market-based incentive program for the federally endangered Golden-cheeked Warbler (Dendroica chrysoparia) was implemented on a trial basis at Fort Hood, an Army training post in Texas, USA. Under this program, recovery credits accumulated by Fort Hood through contracts with private landowners are used to offset unintentional loss of breeding habitat of Golden-cheeked Warblers within the installation. Critical to successful implementation of such programs is the ability to value, in terms of changes to overall species viability, both habitat loss and habitat restoration or protection. In this study, we sought to answer two fundamental questions: Given the same amount of change in breeding habitat, does the change in some patches have a greater effect on metapopulation persistence than others? And if so, can characteristics of a patch (e.g., size or spatial location) be used to predict how the metapopulation will respond to these changes? To answer these questions, we describe an approach for using sensitivity analysis of a metapopulation projection model to predict how changes to specific habitat patches would affect species viability. We used a stochastic, discrete-time projection model based on stage-specific estimates of survival and fecundity, as well as various assumptions about dispersal among populations. To assess a particular patch's leverage, we quantified how much metapopulation viability was expected to change in response to changing the size of that patch. We then related original patch size and distance from the largest patch to each patch's leverage to determine if general patch characteristics could be used to develop guidelines for valuing changes to patches within a metapopulation. We found that both the characteristic that best predicted patch leverage and the magnitude of the relationship changed under different model scenarios. Thus, we were unable to find a consistent set of relationships, and therefore we emphasize the dangers in relying on general guidelines to assess patch value. Instead, we provide an approach that can be used to quantitatively evaluate patch value and identify critical needs for future research.     

Effectiveness of Corridors Relative to Enlargement of Habitat Patches

Abstract:  The establishment of biological corridors between two otherwise isolated habitat patches is a common yet contentious strategy for conserving populations in fragmented landscapes. We compared the effectiveness of corridors with the effectiveness of an alternate conservation strategy, the enlargement of existing habitat patches. We used a spatially explicit population model that simulated population size in two kinds of patches. One patch had a corridor that connected it to a larger "source" patch and the other patch was unconnected and enlarged at the periphery by an area the same size as the corridor. Patch isolation, corridor width, patch size, and the probability that individuals would cross the border from habitat to matrix were varied independently. In general, population size was greater in enlarged patches than in connected patches when patches were relatively large and isolated. Corridor width and the probability of crossing the border from habitat to matrix did not affect the relative benefit of corridors versus patch enlargement. Although biological corridors may mitigate potential effects of inbreeding depression at long time scales, our results suggest that they are not always the best method of conserving fragmented populations.     

Populations in small, ephemeral habitat patches may drive dynamics in a Daphnia magna metapopulation

Migration is the key process to understand the dynamics and persistence of a metapopulation. Many metapopulation models assume a positive correlation between habitat patch size or stability and the number of emigrants. However, few empirical data exist, and habitat patch size and habitat stability may affect dispersal differently than they affect local persistence. Here, we studied the production of the migration stage (i.e., resting eggs called ephippia) of the cladoceran Daphnia magna in a metapopulation consisting of 530 rock pool habitat patches over 25 years. Earlier, the functioning of this metapopulation was explained with a Levins-type metapopulation model or with a mainland-island metapopulation model, based on local extinction and colonization data or time series data, respectively. We used pool volume, hydroperiod length, and number of desiccation events to calculate per-pool production of ephippia (i.e., migration stages). We estimated that populations in small and ephemeral habitat patches produced more than half of the 250 000 to 1 million ephippia that were produced in the metapopulation as a whole per year between 1982 and 2006. Furthermore, these small populations contributed approximately 90% of the ephippia exposed during desiccation events, while the contribution of the long-lived populations in large pools was minimal. We term this an "inverse mainland-island" type metapopulation and propose that populations in small, ephemeral habitat patches may also be the driving force for metapopulation dynamics in other systems.     

Potential Misuse of Avian Density as a Conservation Metric

Abstract: Effective conservation metrics are needed to evaluate the success of management in a rapidly changing world. Reproductive rates and densities of breeding birds (as a surrogate for reproductive rate) have been used to indicate the quality of avian breeding habitat, but the underlying assumptions of these metrics rarely have been examined. When birds are attracted to breeding areas in part by the presence of conspecifics and when breeding in groups influences predation rates, the effectiveness of density and reproductive rate as indicators of habitat quality is reduced. It is beneficial to clearly distinguish between individual‐ and population‐level processes when evaluating habitat quality. We use the term reproductive rate to refer to both levels and further distinguish among levels by using the terms per capita fecundity (number of female offspring per female per year, individual level) and population growth rate (the product of density and per capita fecundity, population level). We predicted how density and reproductive rate interact over time under density‐independent and density‐dependent scenarios, assuming the ideal free distribution model of how birds settle in breeding habitats. We predicted population density of small populations would be correlated positively with both per capita fecundity and population growth rate due to the Allee effect. For populations in the density‐dependent growth phase, we predicted no relation between density and per capita fecundity (because individuals in all patches will equilibrate to the same success rate) and a positive relation between density and population growth rate. Several ecological theories collectively suggest that positive correlations between density and per capita fecundity would be difficult to detect. We constructed a decision tree to guide interpretation of positive, neutral, nonlinear, and negative relations between density and reproductive rates at individual and population levels.     

Dispersal in Spatially Explicit Population Models

Abstract: Ruckelshaus et al. (1997) outlined a simulation model of dispersal between patches in a fragmented landscape. They showed that dispersal success—the proportion of dispersers successfully locating a patch—was particularly sensitive to errors in dispersal mortality and concluded that this limits the utility of spatially explicit population models in conservation biology. I contend that, although they explored error propagation in a simple dispersal model, they did not explore how errors are propagated in spatially explicit population models, as no consideration of population processes was included. I developed a simple simulation model to investigate the effect of varying dispersal success on predictions of patch occupancy and population viability, the conventional outputs of spatially explicit population models. The model simulates births and deaths within habitat patches and dispersal as the transfer of individuals between them. Model predictions were sensitive to changes in dispersal success across a restricted range of within-patch growth rates, which depended on the dispersal initiation mechanism, patch carrying capacities, and number of generations simulated. Predictions of persistence and patch occupancy were generally more sensitive to changes in dispersal success (1) under presaturation rather than saturation dispersal; (2) at lower patch carrying capacities; and (3) over longer time periods. The framework I present provides a means of assessing, quantitatively, the regions of parameter space for which differences in dispersal success are likely to have a large effect on population model outputs. Investigating the effect of the representation of dispersal behavior within the demographic and landscape context provides a more useful assessment of whether our lack of knowledge is likely to cause unacceptable uncertainty in the predictions of spatially explicit population models.     

Should heterogeneity be the basis for conservation? Grassland bird response to fire and grazing.

In tallgrass prairie, disturbances such as grazing and fire can generate patchiness across the landscape, contributing to a shifting mosaic that presumably enhances biodiversity. Grassland birds evolved within the context of this shifting mosaic, with some species restricted to one or two patch types created under spatially and temporally distinct disturbance regimes. Thus, management-driven reductions in heterogeneity may be partly responsible for declines in numbers of grassland birds. We experimentally altered spatial heterogeneity of vegetation structure within a tallgrass prairie by varying the spatial and temporal extent of fire and by allowing grazing animals to move freely among burned and unburned patches (patch treatment). We contrasted this disturbance regime with traditional agricultural management of the region that promotes homogeneity (traditional treatment). We monitored grassland bird abundance during the breeding seasons of 2001-2003 to determine the influence of altered spatial heterogeneity on the grassland bird community. Focal disturbances of patch burning and grazing that shifted through the landscape over several years resulted in a more heterogeneous pattern of vegetation than uniform application of fire and grazing. Greater spatial heterogeneity in vegetation provided greater variability in the grassland bird community. Some bird species occurred in greatest abundance within focally disturbed patches, while others occurred in relatively undisturbed patches in our patch treatment. Henslow's Sparrow, a declining species, occurred only within the patch treatment. Upland Sandpiper and some other species were more abundant on recently disturbed patches within the same treatment. The patch burn treatment created the entire gradient of vegetation structure required to maintain a suite of grassland bird species that differ in habitat preferences. Our study demonstrated that increasing spatial and temporal heterogeneity of disturbance in grasslands increases variability in vegetation structure that results in greater variability at higher trophic levels. Thus, management that creates a shifting mosaic using spatially and temporally discrete disturbances in grasslands can be a useful tool in conservation. In the case of North American tallgrass prairie, discrete fires that capitalize on preferential grazing behavior of large ungulates promote a shifting mosaic of habitat types that maintain biodiversity and agricultural productivity.     

Effects of temperature and precipitation on grassland bird nesting success as mediated by patch size

Grassland birds are declining faster than any other bird guild across North America. Shrinking ranges and population declines are attributed to widespread habitat loss and increasingly fragmented landscapes of agriculture and other land uses that are misaligned with grassland bird conservation. Concurrent with habitat loss and degradation, temperate grasslands have been disproportionally affected by climate change relative to most other terrestrial biomes. Distributions of grassland birds often correlate with gradients in climate, but few researchers have explored the consequences of weather on the demography of grassland birds inhabiting a range of grassland fragments. To do so, we modeled the effects of temperature and precipitation on nesting success rates of 12 grassland bird species inhabiting a range of grassland patches across North America (21,000 nests from 81 individual studies). Higher amounts of precipitation in the preceding year were associated with higher nesting success, but wetter conditions during the active breeding season reduced nesting success. Extremely cold or hot conditions during the early breeding season were associated with lower rates of nesting success. The direct and indirect influence of temperature and precipitation on nesting success was moderated by grassland patch size. The positive effects of precipitation in the preceding year on nesting success were strongest in relatively small grassland patches and had little effect in large patches. Conversely, warm temperatures reduced nesting success in small grassland patches but increased nesting success in large patches. Mechanisms underlying these differences may be patch‐size‐induced variation in microclimates and predator activity. Although the exact cause is unclear, large grassland patches, the most common metric of grassland conservation, appears to moderate the effects of weather on grassland‐bird demography and could be an effective component of climate‐change adaptation.     

Blue Tits use fledgling quantity and quality as public information in breeding site choice

Public information (PI), which is the information that can be derived from the behavior and performance of conspecifics, has been demonstrated to be used in many fitness-enhancing decisions. In the context of breeding habitat choice, PI use has been called "habitat copying." We experimentally tested the existence of habitat copying in the Blue Tit (Cyanistes caeruleus), a nonmigratory, short-lived hole-nesting bird. We manipulated the mean number of fledglings raised locally (quantity) and their condition (quality) as components of PI by transferring nestlings from Decreased (D) patches to Increased (I) patches. Our manipulations caused a negative relationship between fledgling quantity and quality that does not exist naturally: I patches had a higher number of fledglings that were in poorer condition, whereas D patches had a lower number in better condition. Control (C) patches, whether manipulated or not, had intermediate levels in terms of fledgling quantity and quality. Adult emigration the following year was higher from D than from C or I patches. Similarly, adult dispersal distance decreased for individuals coming from D to C to I patches. This suggests that resident breeders rely mainly on fledgling quantity to make emigration decisions. Emigration patterns of juveniles did not vary in relation to our patch manipulation. Immigration rates were higher and similar in I and D patches than in C patches. Hence, immigrant Blue Tits seem to rely on one of the manipulated components of PI and are insensitive to the discrepancy between fledgling quantity and quality. This shows that even nonmigratory species, such as Blue Tits, may use PI in their dispersal decisions but weigh its components differently for emigration and immigration. Differences among species in the importance of PI in breeding habitat choices may be explained by differences in life histories.     

Using network centrality measures to manage landscape connectivity

We use a graph-theoretical landscape modeling approach to investigate how to identify central patches in the landscape as well as how these central patches influence (1) organism movement within the local neighborhood and (2) the dispersal of organisms beyond the local neighborhood. Organism movements were theoretically estimated based on the spatial configuration of the habitat patches in the studied landscape. We find that centrality depends on the way the graph-theoretical model of habitat patches is constructed, although even the simplest network representation, not taking strength and directionality of potential organisms flows into account, still provides a coarse-grained assessment of the most important patches according to their contribution to landscape connectivity. Moreover, we identify (at least) two general classes of centrality. One accounts for the local flow of organisms in the neighborhood of a patch, and the other accounts for the ability to maintain connectivity beyond the scale of the local neighborhood. Finally, we study how habitat patches with high scores on different network centrality measures are distributed in a fragmented agricultural landscape in Madagascar. Results show that patches with high degree and betweenness centrality are widely spread, while patches with high subgraph and closeness centrality are clumped together in dense clusters. This finding may enable multispecies analyses of single-species network models.     

Inferring Metapopulation Dynamics from Patch-Level Incidence of Florida Scrub Plants

Spatial structure and dynamics of multiple populations may explain species distribution patterns in patchy communities with heterogeneous disturbance regimes, especially when species have poor dispersal. The endemic-rich Florida (U.S.A.) rosemary scrub occupies about 4% of the west portion of Archbold Biological Station and occurs scattered within a matrix of less xeric vegetation. Longer fire-return times and higher frequency of open patches in rosemary scrub provide favorable habitat for many plant species. Occupancy of 123 species of vascular plants and ground lichens in 89 patches was determined by repeated site surveys. About two-thirds of the species occurring at more than 14 patches had a significant logistic regression of presence on time-since-fire, patch size, patch isolation, or their interactions. Species with presence related to the interaction between patch isolation and patch size were primarily herbs and small shrubs specializing in rosemary scrub. These results suggest the importance of spatial characteristics of the landscape for population turnover of these species. An incidence-based metapopulation model was used to predict extinction and colonization probabilities of those species with presence in rosemary scrub patches related to the studied spatial variables. This is the first attempt to apply incidence-based metapopulation models to plants. The results showed stronger effects of patch size and patch isolation on extinction probabilities of herbs than on those of woody species. Because of their effect on spatial heterogeneity and habitat availability, fire suppression and habitat destruction may decrease persistence probabilities for these rosemary scrub specialists, many of which are endangered species.     

Conservation of Fragmented Populations

In this paper we argue that landscape spatial structure is of central importance in understanding the effects of fragmentation on population survival. Landscape spatial structure is the spatial relationships among habitat patches and the matrix in which they are embedded. Many general models of subdivided populations make the assumptions that (1) all habitat patches are equivalent in size and quality and (2) all local populations (in the patches) are equally accessible by dispersers. Models that gloss over spatial details of landscape structure can be useful for theoretical developments but will almost always be misleading when applied to real-world conservation problems. We show that local extinctions of fragmented populations are common. From this it follows that recolonization of local extinctions is critical for regional survival of fragmented populations. The probability of recolonization depends on (1) spatial relationships among landscape elements used by the population, including habitat patches for breeding and elements of the inter-patch matrix through which dispersers move, (2) dispersal characteristics of the organism of interest, and (3) temporal changes in the landscape structure. For endangered species, which are typically restricted in their dispersal range and in the kinds of habitat through which they can disperse, these factors are of primary importance and must be explicitly considered in management decisions.     

Evaluating the effect of corridors and landscape heterogeneity on dispersal probability: a comparison of three spatially explicit modelling approaches

Spatially explicit simulation models of varying degree of complexity are increasingly used in landscape and species management and conservation. The choice as to which type of model to employ in a particular situation, is however, far too often governed by logistic constraints and the personal preferences of the modeller, rather than by a critical evaluation of model performance. We present a comparison of three common spatial simulation approaches (patch-based incidence-function model (IFM), individual-based movement model (IBMM), individual-based population model including detailed behaviour and demographics (IBPM)). The IBPM was analysed in two versions (IBPM_st and IBPM_dyn). Both assumed spatial heterogeneity of the matrix, but the IBPM_dyn in addition included temporal matrix dynamics. The models were developed with a shared minimum objective, namely to predict dynamics of individuals or populations in space given a specific configuration of habitat patches. We evaluated how the choice of model influenced predictions regarding the effect of patch and corridor configuration on dispersal probabilities and the number of successful immigrants of a simulated small mammal. Model results were analysed both at the level of the entire habitat network and at the level of individual patches.All models produced similar rankings of alternative habitat networks, but large discrepancies existed between absolute estimates of dispersal probabilities and the number of successful immigrants predicted by the different models. Generally, predicted dispersal probabilities were highest in the IBMM, intermediate in the IFM and the IBPM_st and lowest in the IBPM_dyn. Observed differences were due both to differences in implementation (e.g. raster versus vector-based movement algorithms), the chosen level of detail in landscape representation (e.g. matrix complexity) and the degree of behavioural realism included in the models (e.g. demography, differentiated mortality).The advantages and disadvantages of the three modelling approaches are discussed, as are the implications of the results for the recommended use of the three types of models in practical management.     

A model for behavioral regulation of metapopulation dynamics

Habitat fragmentation can decrease local population persistence by reducing connectivity, which is a function of dispersal of individuals among habitat fragments. Dispersal is often treated as diffusion in population models, even though for many species it is a result of a series of behavioral decisions. We developed a metapopulation model to explore the potential importance of dispersal behaviors in driving metapopulation dynamics. We incorporated types of behavior that affect dispersal—colonization inhibiting, colonization enhancing, extinction inhibiting, extinction enhancing, rescue enhancing, rescue inhibiting—into Levins’ (1969) metapopulation model and projected occupancy rates for a variety of parameter values. Examples from the literature of behaviors associated with each of these parameters are provided. Our model simplifies into previously published metapopulation models that incorporate only a single behavior, and we present a density-dependent rescue function that leads to multiple non-zero equilibria. We found a variety of behavioral effects on metapopulations. Rescue enhancement fills patches faster than does colonization enhancement or extinction inhibition, and declines in patch occupancy are moderate with extinction enhancement, but colonization inhibition causes metapopulation extinction. We also found that with colonization and extinction inhibitions, equilibrium patch occupancy is inversely related to patch turnover rate. With density-dependent rescue, persistence depends not only on the strength of the strong rescue effect, but also on having a sufficient initial fraction of patches occupied; the stronger the rescue effect, the lower this fraction can be. This study suggests that dispersal behavior can have strong influences on metapopulation dynamics. It confirms the importance of understanding the relationship between landscape structure and dispersal behavior in understanding population persistence.     

Possible causes and consequences of philopatry and breeding dispersal in red-backed shrikes <Emphasis Type="BoldItalic">Lanius collurio</Emphasis>

Studies of animal breeding dispersal have often focused on possible causes, whereas its adaptive significance has received less attention. Using an information-theoretic approach, we assessed predictions of four hypotheses relating to causes and consequences of breeding dispersal in a migratory passerine, the red-backed shrike Lanius collurio. As predicted by the reproductive performance hypothesis, probability of breeding dispersal in females (though not in males) decreased with increasing annual average number of fledglings produced in the past year, but there was no association with conspecific reproductive performance in either sex. The site choice hypothesis, stating that individuals disperse to improve breeding site quality, received support in males only, as dispersal probability was positively associated to a measure indicating low territory quality. The social constraints hypothesis, referring to dispersal in relation to intraspecific interactions, received little support in either sex. The predation risk hypothesis was hardly supported either. Consequences of dispersal were marginal in both sexes because neither fledgling production in females, nor territory quality in males improved after dispersal. In addition, males settled on territories closer to the forest edge than those occupied predispersal, which is opposite to the prediction of the predation risk hypothesis. We conclude that own reproductive success was the major factor determining dispersal behavior in females, whereas territory quality and possibly predation risk were most important in males. Overall, breeding dispersal appeared not to be adaptive in this dense population inhabiting an optimal habitat.