Landscape structure shapes habitat finding ability in a butterfly

Land-use intensification and habitat fragmentation is predicted to impact on the search strategies animals use to find habitat. We compared the habitat finding ability between populations of the speckled wood butterfly (Pararge aegeria L.) from landscapes that differ in degree of habitat fragmentation. Naïve butterflies reared under standardized laboratory conditions but originating from either fragmented agricultural landscapes or more continuous forested landscapes were released in the field, at fixed distances from a target habitat patch, and their flight paths were recorded. Butterflies originating from fragmented agricultural landscapes were better able to find a woodlot habitat from a distance compared to conspecifics from continuous forested landscapes. To manipulate the access to olfactory information, a subset of individuals from both landscape types were included in an antennae removal experiment. This confirmed the longer perceptual range for butterflies from agricultural landscapes and indicated the significance of both visual and olfactory information for orientation towards habitat. Our results are consistent with selection for increased perceptual range in fragmented landscapes to reduce dispersal costs. An increased perceptual range will alter the functional connectivity and thereby the chances for population persistence for the same level of structural connectivity in a fragmented landscape.     

Does the type of matrix matter? A quantitative review of the evidence

It has been increasingly recognized that the type of matrix surrounding habitat patches can affect biodiversity in landscapes, but there were only qualitative reviews of the subject focused on particular taxonomic groups. We present a quantitative review of studies from 1985 to 2008 that compared effects of different matrix types on individuals, populations and communities. We compiled 104 studies, most on animals, covering a broad range of landscape types and spatial scales. Most studies were empirical, focused on individuals and communities, and evaluated abundance/richness in the patch as the dependent variable. The type of matrix surrounding habitat patches influenced the studied parameters in 95% of the studies, but such effects were overall smaller compared to patch size or isolation effects. Matrix type effects were strongly species-specific, with different species responding differently to matrix type in 96% of studies comparing species or group of species. In 88% of studies, matrix types more similar in structure to the patch had higher quality for the studied organisms from the point of view of functional connectivity. Overall, the type of matrix is important, but patch size and isolation are the main determinants of ecological parameters in landscapes. Matrix quality generally increases with increasing structural similarity with habitat patches, a pattern that could be used as a general guideline for management of the matrix in fragmented landscapes.     

Between‐patch natal dispersal declines with increasing natal patch size and distance to other patches in the endangered Southern Dunlin Calidris alpina schinzii

Natal dispersal has profound consequences for populations through the movement of individuals and genes. Habitat fragmentation reduces structural connectivity by decreasing patch size and increasing isolation, but understanding of how this impacts dispersal and the functional connectivity of landscapes is limited because many studies are constrained by the size of the study areas or sample sizes to accurately capture natal dispersal. We quantified natal dispersal probability and natal dispersal distances in a small migratory shorebird, the Southern Dunlin Calidris alpina schinzii, with data from two extensively monitored endangered metapopulations breeding in Sweden and Finland. In both metapopulations philopatry was strong, with individuals returning to or close to their natal patches more often than expected by chance, consistent with the patchy distribution of their breeding habitat. Dispersal probabilities were lower and dispersal distances were shorter in Sweden. These results provide a plausible explanation for the observed inbreeding and population decline of the Swedish population. The differences between Sweden and Finland were explained by patch‐specific differences. Between‐patch dispersal decreased with increasing natal patch size and distance to other patches. Our results suggest that reduced connectivity reduces movements of the philopatric Dunlin, making it vulnerable to the effects of inbreeding. Increasing connectivity between patches should thus be one of the main goals when planning future management. This may be facilitated by creating a network of suitably sized patches (20–100 ha), no more than 20 km apart from each other, from existing active patches that may work as stepping stones for movement, and by increasing nest success and pre‐fledging survival in small patches.     

How do birds search for breeding areas at the landscape level? Interpatch movements of male ortolan buntings

Animal movements at large spatial scales are of great importance in population ecology, yet little is known due to practical problems following individuals across landscapes. We studied the whole Norwegian population of a small songbird (ortolan bunting, Emberiza hortulana ) occupying habitat patches dispersed over nearly 500 km². Movements of colour-ringed males were monitored during ten years, and extensive long-distance dispersal was recorded. More than half of all cases of breeding dispersal took place within one breeding season, and males moved up to 43 km between singing territories, using 1–22 d. Natal dispersal was usually to a habitat patch close to the natal patch, or within the natal patch if it was large. Breeding dispersal movements were often long-distance, beyond neighbouring patches, and up to 11–19 patches were overflown. Movements of at least 6–9 km across areas of unsuitable habitat occurred regularly. The number of patches visited was low (1–4) even though search costs in terms of time spent moving from one site to another were relatively low (often only a few days even for distances >10 km). Most males seemed to use a threshold tactic when choosing a patch, but returns to previously visited patches were recorded, including some cases of commuting. In conclusion, male ortolan buntings have a surprising ability to move quickly at the landscape level, and this resulted in a high connectivity of patches. We discuss our results in relation to optimal searching strategies, in particular the use of within-breeding season versus post-breeding season search, conspecific attraction and adaptive late arrival of young birds.     

The taste of nectar – a neglected area of pollination ecology

We investigated the impact of landscape structure on landscape connectivity using a combination of simulation and empirical experiments. In a previous study we documented the movement behaviour of a specialized goldenrod beetle ( Trirhabda borealis Blake) in three kinds of patches: habitat (goldenrod) patches and two types of matrix patch (cut vegetation and cut vegetation containing camouflage netting as an impediment to movement). In the current study, we used this information to construct simulation and experimental landscapes consisting of mosaics of these three patch types, to study the effect of landscape structure on landscape connectivity, using the T. borealis beetle as a model system. In the simulation studies, landscape connectivity was based on movements of individual beetles, and was measured in six different ways. The simulations revealed that the six measures of landscape connectivity were influenced by different aspects of landscape structure, suggesting that: (1) landscape connectivity is a poorly defined concept, and (2) the same landscape may have different landscape connectivity values when different measures of landscape connectivity are used. There were two general predictions that held over all measures of landscape connectivity: (1) increasing interpatch distance significantly decreased landscape connectivity and (2) the influence of matrix elements on landscape connectivity was small in comparison to the influence of habitat elements. Empirical mark-release-resight experiments using Trirhabda beetles in experimental landscapes supported the simulation results.     

Modelling the third dimension: Incorporating topography into the movement rules of an individual-based spatially explicit population model

A wide variety of topographical and environmental elements have been shown or proposed to influence the movement decisions of dispersing animals. Most real landscapes have topographical elements such as hills, valleys and urban developments, which can all act to modify a species’ perceptual range and directly influence movement behaviour. If a visual-based perceptual ability enables a dispersing individual to locate suitable habitat patches at a distance, then it is to be expected that topographical features would act to modify the overall success of this strategy. However, the majority of individual-based Spatially Explicit Population Models (SEPM) employ only two-dimensional landscapes.To investigate the effects of topographical elevation on dispersal patterns, a three-dimensional visual-based perceptual range algorithm was added to the dispersal rules of an individual-based SEPM. To explore the possible influences of a behavioural-based response to topography, an algorithm modelling valley-seeking behaviour was also developed. The performance of both algorithms was compared with that of a two-dimensional visual-based perceptual range algorithm. The overall consequences of dispersal under each algorithm were measured by recording population sizes in a target wood in the centre of a modelled, real landscape.The size of the population in the target wood, modelled using both of the three-dimensional algorithms, exhibited sensitivity to the direction of dispersal in interaction with perceptual range, which differed from that predicted by the two-dimensional approach. Population size was dependant on the spatial configuration of habitat patches and on the topography of the landscape, both of which could guide dispersers either towards or away from the target patch depending on the particular combinations of dispersal directions and perceptual ranges selected. Topography was found to have a greater effect on dispersal at shorter perceptual ranges, and thresholds in the results for all three algorithms suggested the existence of species and landscape dependant optimal perceptual ranges. It is recommended that both topography and topographical-based dispersal-altering algorithms, commensurate with the studied species’ behaviour, be incorporated into the movement rule-base of dispersal simulation models. The modelling of topography and its effects on movement in patchy landscapes are seen as essential ingredients in future landscape planning.     

Generalizing matrix structure affects the identification of least-cost paths and patch connectivity

Understanding and assessing landscape connectivity is often a primary goal when studying patchy or spatially structured populations. It is commonly accepted that the matrix plays a role in determining connectivity; however, it is not clear how the process of assessing connectivity is impacted by different ways in which the matrix may be represented, particularly if matrix structure is generalized to expedite analysis. We conducted a controlled experiment using computer simulations to evaluate the impact of increasing levels of matrix generalization on connectivity assessment using a constant arrangement of habitat patches. We varied matrix generalization for six simulated landscape patterns by adjusting the number of classes and level of pattern grain in 16 ways to yield sets of landscapes for which the matrix ranged from not generalized (i.e., heterogeneous) to completely generalized (i.e., homogeneous) while habitat placement was held constant. Least-cost paths were calculated for each landscape, and a spatial interaction model (SIM) was implemented to model the counts of patch inflows (immigration) and pairwise exchange. Applying a SIM allowed us to generate absolute outputs and explicitly compare the effect of changes to matrix generalization on connectivity. We found that both assessment of connections (i.e., measured distance and spatial delineation of least-cost paths) and patch inflows/pairwise exchange were highly sensitive to matrix generalization and that effects were inconsistent and unpredictable across the range of matrix representations, especially when estimating connectivity for individual patches. We conclude that matrix pattern may have an underappreciated effect on connectivity and that least-cost path delineation and connectivity assessment may be very sensitive to generalizations of the matrix. We suggest that sensitivity analysis of the matrix representation should be performed when conducting connectivity analyses.     

Limited perceptual range and anemotaxis in marsh rice ratsOryzomys palustris

Dispersal among suitable habitat patches could be crucial to the regional persistence of wetland species. Dispersal success of animals will depend, in part, on the distance from which they can detect suitable habitat (ie, perceptual range). We investigated the perceptual range of marsh rice rats Oryzomys palustris Harlan, 1837 during the wet season in a region of central Florida characterized by small, isolated depression marshes. O. palustris had a limited perceptual range (<10 m) indicating that individuals moving through unfamiliar habitat have restricted information from which to make movement decisions. O. palustris displayed anemotaxis during our experiment; individuals generally moved either upwind or downwind but rarely crosswind. This anemotaxis might reflect use of wind by rice rats to maintain a straight course while searching for new habitat located beyond their perceptual range.     

Conspecific and heterospecific attraction in assessments of functional connectivity

Functional connectivity is known to have an important, positive influence on species persistence. Measurements of functional connectivity traditionally focus on structural attributes of landscapes such as the distance and matrix type between habitat patches as well as on how species interact with those structural attributes. However, we propose that the social behavior of a species, through conspecific and heterospecific attraction, will also impact connectivity by changing how dispersers move with respect to each other and occupied patches. We analyzed functional connectivity patterns using circuit and graph theory for golden-headed lion tamarins (Leontopithecus chrysomelas) in Brazil under three scenarios. In the first scenario, we looked at connectivity without the effects of attraction under varying maximum dispersal distance and ecological movement cost thresholds. In the second scenario, we allowed dispersers to travel over more hostile matrix than they normally would to reach an occupied patch. In the final scenario, we allowed dispersers to move only to occupied patches. We found that, according to the first scenario, range-wide functional landscape connectivity for golden-headed lion tamarins is low at realistic maximum dispersal distance and movement cost thresholds. Incorporating the effects of conspecific or heterospecific attraction would increase functional connectivity, in the case of scenario two, or decrease functional connectivity, in the case of scenario three. Because conspecific/heterospecific attraction can have an impact on movement for some species, this factor should be incorporated in assessments of functional connectivity patterns for social species and others where patch occupancy is likely to influence the movements of dispersers.     

Adaptive Patch Searching Strategies in Fragmented Landscapes

The search strategies dispersers employ to search for new habitat patches affect individuals’ search success and subsequently landscape connectivity and metapopulation viability. Some evidence indicates that individuals within the same species may display a variety of behavioural patch searching strategies rather than one species-specific strategy. This may result from landscape heterogeneity. We modelled the evolution of individual patch searching strategies in different landscapes. Specifically, we analysed whether evolution can favour different, co-existing, behavioural search strategies within one population and to what extent this coexistence of multiple strategies was dependent on landscape configuration. Using an individual-based simulation model, we studied the evolution of patch searching strategies in three different landscape configurations: uniform, random and clumped. We found that landscape configuration strongly influenced the evolved search strategy. In uniform landscapes, one fixed search strategy evolved for the entire spatially structured population, while in random and clumped landscapes, a set of different search strategies emerged. The coexistence of several search strategies also strongly depended on the dispersal mortality. We show that our result can affect landscape connectivity and metapopulation dynamics. Co-ordinating editor: N. Yamamura     

The matrix matters: effective isolation in fragmented landscapes

Traditional approaches to the study of fragmented landscapes invoke an island-ocean model and assume that the nonhabitat matrix surrounding remnant patches is uniform. Patch isolation, a crucial parameter to the predictions of island biogeography and metapopulation theories, is measured by distance alone. To test whether the type of interpatch matrix can contribute significantly to patch isolation, I conducted a mark-recapture study on a butterfly community inhabiting meadows in a naturally patchy landscape. I used maximum likelihood to estimate the relative resistances of the two major matrix types (willow thicket and conifer forest) to butterfly movement between meadow patches. For four of the six butterfly taxa (subfamilies or tribes) studied, conifer was 3-12 times more resistant than willow. For the two remaining taxa (the most vagile and least vagile in the community), resistance estimates for willow and conifer were not significantly different, indicating that responses to matrix differ even among closely related species. These results suggest that the surrounding matrix can significantly influence the "effective isolation" of habitat patches, rendering them more or less isolated than simple distance or classic models would indicate. Modification of the matrix may provide opportunities for reducing patch isolation and thus the extinction risk of populations in fragmented landscapes.     

Identifying multiscale habitat factors influencing koala (Phascolarctos cinereus) occurrence and management in Ballarat, Victoria, Australia

Summary   Modelling for the conservation of koala ( Phascolarctos cinereus ) populations has primarily focused on natural habitat variables (e.g. tree species, soil types and soil moisture). Until recently, limited consideration has been given to modelling the effects of the landscape context (e.g. habitat area, habitat configuration and roads). Yet, the combined influence of natural habitats and anthropogenic impacts at multiple spatial scales are likely to be important determinants of where koala populations occur and remain viable in human-modified landscapes. The study tested the importance of multiscale habitat variables on koala occurrence in Ballarat, Victoria, Australia. The models focused at three spatial scales: site ( <  1 ha), patch (1–100 ha), and landscape (100–1000 s ha). Logistic regression and hierarchical partitioning analyses were used to rank alternative models and key explanatory variables.
The results showed that an increased likelihood of koala presence in fragmented landscapes in the urban–forest interface (as opposed to larger blocks of forest habitat) can best be explained by the positive effects of soil fertility and the presence of preferred koala tree species in these fragmented areas. If koalas are to be effectively conserved in Ballarat, it is critical to (i) protect remaining core areas of high-quality habitat, including regenerating areas; (ii) protect scattered habitat patches which provide connectivity; and (iii) develop and implement habitat restoration programmes to improve habitat connectivity and enhance opportunities for safe koala movement between habitat patches intersected by main roads.     

The importance of forest patch networks for the conservation of the Thorn-tailed Rayaditos in central Chile

Conservation of forest birds in fragmented landscapes requires not only determining the critical patch characteristics influencing local population persistence but also identifying patch networks providing connectivity and suitable habitat conditions necessary to ensure regional persistence. In this study, we assessed the importance of patch attributes, patch connectivity, and network components (i.e., groups of interconnected patches) in explaining the occupancy pattern of the Thorn-tailed Rayadito (Aphrastura spinicauda), a forest bird species of central Chile. Using a daily movement threshold distance, we identified a total of 16 network components of sclerophyllous forest within the study area. Among those components, patch area and vegetation structure-composition were important predictors of patch occupancy. However, the inclusion of patch connectivity and component size (i.e., the area of a network component) into the models greatly increases the models’ accuracy and parsimony. Using the best-fitted model, a total of 33 patches were predicted to be occupied by rayaditos within the study area, but such occupied patches were distributed in only six network components. These results suggest that persistence of rayaditos in central Chile requires the maintenance of large single patches and patch networks providing habitat and connectivity.     

Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.     

Gene flow and pathogen transmission among bobcats (Lynx rufus) in a fragmented urban landscape

Urbanization can result in the fragmentation of once contiguous natural landscapes into a patchy habitat interspersed within a growing urban matrix. Animals living in fragmented landscapes often have reduced movement among habitat patches because of avoidance of intervening human development, which potentially leads to both reduced gene flow and pathogen transmission between patches. Mammalian carnivores with large home ranges, such as bobcats (Lynx rufus), may be particularly sensitive to habitat fragmentation. We performed genetic analyses on bobcats and their directly transmitted viral pathogen, feline immunodeficiency virus (FIV), to investigate the effects of urbanization on bobcat movement. We predicted that urban development, including major freeways, would limit bobcat movement and result in genetically structured host and pathogen populations. We analysed molecular markers from 106 bobcats and 19 FIV isolates from seropositive animals in urban southern California. Our findings indicate that reduced gene flow between two primary habitat patches has resulted in genetically distinct bobcat subpopulations separated by urban development including a major highway. However, the distribution of genetic diversity among FIV isolates determined through phylogenetic analyses indicates that pathogen genotypes are less spatially structured-exhibiting a more even distribution between habitat fragments. We conclude that the types of movement and contact sufficient for disease transmission occur with enough frequency to preclude structuring among the viral population, but that the bobcat population is structured owing to low levels of effective bobcat migration resulting in gene flow. We illustrate the utility in using multiple molecular markers that differentially detect movement and gene flow between subpopulations when assessing connectivity.     

Patch occupancy by squirrels in fragmented deciduous forest: effects of behavior

We tested whether species-specific behavioral traits could explain patterns of habitat patch occupancy by five different squirrel species in Ontario, Canada: the northern and southern flying squirrel (Glaucomys sabrinus and Glaucomys volans), the North American red squirrel (Tamiasciurus hudsonicus), the eastern chipmunk (Tamias striatus), and the eastern gray squirrel (Sciurus carolinensis). Flying squirrel species exhibit group nesting in winter, which may put them at risk of extirpation in small patches with few individuals to contribute to group nests. Flying squirrels are also volant, potentially making non-treed matrix a barrier. Our surveys revealed that G. sabrinus was most likely to occur in large patches that were embedded in landscapes with low connectivity, and least likely to occur in small patches in highly connected landscapes. Conversely, G. volans was most likely to occur in large, well-connected patches and least likely to occur in small, unconnected patches. Patch occupancy by the cursorial squirrels was not strongly influenced by patch area or isolation. These findings reinforce previous studies suggesting that an understanding of species-specific traits such as behavior is an important consideration when interpreting habitat fragmentation effects.     

Area-dependent migration by ringlet butterflies generates a mixture of patchy population and metapopulation attributes

Interpretation of spatially structured population systems is critically dependent on levels of migration between habitat patches. If there is considerable movement, with each individual visiting several patches, there is one ”patchy population”; if there is intermediate movement, with most individuals staying within their natal patch, there is a metapopulation; and if (virtually) no movement occurs, then the populations are separate (Harrison 1991, 1994). These population types actually represent points along a continuum of much to no mobility in relation to patch structure. Therefore, interpretation of the effects of spatial structure on the dynamics of a population system must be accompanied by information on mobility. We use empirical data on movements by ringlet butterflies, Aphantopus hyperantus, to investigate two key issues that need to be resolved in spatially-structured population systems. First, do local habitat patches contain largely independent local populations (the unit of a metapopulation), or merely aggregations of adult butterflies (as in patchy populations)? Second, what are the effects of patch area on migration in and out of the patches, since patch area varies considerably within most real population systems, and because human landscape modification usually results in changes in habitat patch sizes? Mark-release-recapture (MRR) data from two spatially structured study systems showed that 63% and 79% of recaptures remained in the same patch, and thus it seems reasonable to call both systems metapopulations, with some capacity for separate local dynamics to take place in different local patches. Per capita immigration and emigration rates declined with increasing patch area, while the resident fraction increased. Actual numbers of emigrants either stayed the same or increased with area. The effect of patch area on movement of individuals in the system are exactly what we would have expected if A. hyperantus were responding to habitat geometry. Large patches acted as local populations (metapopulation units) and small patches simply as locations with aggregations (units of patchy populations), all within 0.5 km². Perhaps not unusually, our study system appears to contain a mixture of metapopulation and patchy-population attributes.     

Using connectivity metrics in conservation planning – when does habitat quality matter?

Aim The objective of conservation planning is often to prioritize patches based on their estimated contribution to metapopulation or metacommunity viability. The contribution that an individual patch makes will depend on its intrinsic characteristics, such as habitat quality, as well as its location relative to other patches, its connectivity. Here we systematically evaluate five patch value metrics to determine the importance of including an estimate of habitat quality into the metrics. Location We tested the metrics in landscapes designed to represent different degrees of variability in patch quality and different levels of patch aggregation. Methods In each landscape, we simulated population dynamics using a spatially explicit, continuous time metapopulation model linked to within patch logistic growth models. We tested five metrics that are used to estimate the contribution that a patch makes to metapopulation viability: two versions of the probability of connectivity index, two versions of patch centrality (a graph theory metric) and the metapopulation capacity metric. Results All metrics performed best in environments where patch quality was very variable and high quality patches were aggregated. Metrics that incorporated some measure of patch quality did better in all environments, but did particularly well in environments with high variance of patch quality and spatial aggregation of good quality patches. Main conclusions Including an estimate of patch quality significantly increased the ability of a given connectivity metric to rank correctly habitat patches according to their contribution to metapopulation viability. Incorporating patch quality is particularly important in landscapes where habitat quality is highly variable and good quality patches are spatially aggregated. However, caution should be used when applying patch metrics to homogeneous landscapes, even if good estimates of patch quality are available. Our results demonstrate that landscape structure and the degree of variability in patch quality need to be assessed prior to selecting a suitable method for estimating patch value.     

Matrix heterogeneity affects population size of the harvest mice: Bayesian estimation of matrix resistance and model validation

It has been increasingly recognized that landscape matrices are an important factor determining patch connectivity and hence the population size of organisms living in highly fragmented landscapes. However, most previous studies estimated the effect of matrix heterogeneity using prior information regarding dispersal or habitat preferences of a focal organism. Here we estimated matrix resistance of harvest mice in agricultural landscapes using a novel pattern‐oriented modeling with Bayesian estimation and no prior information, and then conducted model validation using data sets independent from those used for model construction. First, we investigated the distribution patterns of harvest mice for approximately 400 habitat patches, and estimated matrix resistance for different matrix types using statistical models incorporating patch size, patch environment, and patch connectivity. We used Bayesian estimation with a Markov chain Monte Carlo algorithm, and searched for appropriate matrix resistance that best explained the distribution pattern. Patch connectivity as well as patch quality was an important determinant of local population size for the harvest mice. Moreover, matrix resistance was far from uniform, with rice and crop fields exhibiting low resistance and forests, creeks, roads and residential areas showing much higher resistance. The deviance explained by this model (heterogeneous matrix model) was much larger than that obtained by the model with no consideration of matrix heterogeneity (homogeneous matrix model). Second, we obtained distribution data from five additional landscapes that were more fragmented than that used for model construction, and used them for model validation. The heterogeneous matrix model well predicted the population size for four out of five landscapes. In contrast, the homogeneous model considerably overestimated population sizes in all cases. Our approach is widely applicable to species living in fragmented landscapes, especially those for which prior information regarding movement or dispersal is difficult to obtain.     

Dispersal and egg shortfall in Monarch butterflies: what happens when the matrix is cleaned up?

1. We use an individual‐based model describing the life of a monarch butterfly, which utilises milkweeds both aggregated in patches and scattered across the wider landscape as a substrate for laying eggs. The model simplifies the metapopulation of milkweed habitat patches by representing them as a proportion of the overall landscape, with the rest of the landscape considered matrix, which may contain some low density of milkweed plants. 2. The model simulates the number of eggs laid daily by a butterfly as it searches for hosts. The likelihood of finding hosts is related to the density of plants and the search ability of the butterfly. For an empty matrix, remaining in a habitat patch results in more eggs laid. However individuals that are good searchers have almost equivalent success without remaining in a habitat patch. These individuals are most affected by the presence of hosts in the matrix. 3. Given realistic values of habitat patch availability, our model shows that the presence of plants at a low density in the matrix has a substantial impact on the number of eggs laid; removing these plants can reduce lifetime potential fecundity by ca. 20%. These results have implications for monarch butterflies inhabiting agricultural landscapes, in which genetically modified soybean that is resistant to herbicides has resulted in the decimation of milkweeds over large areas.