Grassland songbird occurrence on remnant prairie patches is primarily determined by landscape characteristics

Context

North American grassland songbird populations have declined significantly due to habitat loss and fragmentation. Understanding the influence of the surrounding landscape on prairie fragment occupancy is vital for predicting the fate of grassland birds in these heavily altered landscapes.

Objectives

We examined the relative importance of local and landscape variables on grassland bird occupancy of prairie fragments using a focal-patch study. We also investigated the spatial scale at which landscape variables were most influential.

Methods

We surveyed birds on 29 unplowed prairie fragments in western Minnesota and eastern North and South Dakota. We quantified local habitat on the fragment using vegetation surveys and aerial photographs and the landscape surrounding the fragment out to 4 km using aerial photographs. We analyzed occupancy using multi-model approaches applied to multiple logistic regression.

Results

Of 38 species encountered, nine were neither too rare nor too abundant to be analyzed. Predictors of patch occupancy were unique for each bird species, yet general patterns emerged. For eight species, landscape variables were more important than local variables. Mostly, those landscape variables measured configuration (e.g., edge density) and not composition (e.g., percent cover of a particular matrix element). Landscape effects were mostly from variables measured at the greatest extents from the prairie fragment.

Conclusions

Using a focal-patch study design we demonstrated the importance of the surrounding landscape, often out to 4 km from the fragment edge, on prairie occupancy by grassland birds. Effective management of grassland songbirds will require attention to the landscape context of prairie fragments.

     

The influence of landscape relative to site and patch variables on primate distributions in the Colombian Llanos

Context

Primates are an important component of biodiversity in tropical regions. However, many studies on the effects of habitat change on primates ignore the relative influence of landscape composition and configuration.

Objectives

This study addresses the question: how important are landscape-scale forest area and composition relative to patch-scale (1–1080 ha) and site-scale (transect of 1 km) habitat variables for the occupancy and abundance of four primate species in the Colombian Llanos.

Methods

Using a randomly stratified survey design, 81 fragments were surveyed for primate occupancy and abundance. We used zero-inflated models to test the relative influence of landscape-scale, patch-scale and site-scale variables on occupancy and abundance for each species. A 95% confidence set of models was constructed using the cumulative Akaike weight for each model and the relative importance of each set of variables calculated for each primate species.

Results

Occupancy was determined by a combination of site-scale, patch-scale and landscape-scale variables but this varied substantially among the primate species.

Conclusion

Our study highlights the importance of managing primates at a range of scales that considers the relative importance of site-, patch- and landscape-scale variables.

     

The plant functional traits that explain species occurrence across fragmented grasslands differ according to patch management,isolation, and wetness

Context

Landscape fragmentation significantly affects species distributions by decreasing the number and connectivity of suitable patches. While researchers have hypothesized that species functional traits could help in predicting species distribution in a landscape, predictions should depend on the type of patches available and on the ability of species to disperse and grow there.

Objectives

To explore whether different traits can explain the frequency of grassland species (number of occupied patches) and/or their occupancy (ratio of occupied to suitable patches) across a variety of patch types within a fragmented landscape.

Methods

We sampled species distributions over 1300 grassland patches in a fragmented landscape of 385 km² in the Czech Republic. Relationships between functional traits and species frequency and occupancy were tested across all patches in the landscape, as well as within patches that shared similar management, wetness, and isolation.

Results

Although some traits predicting species frequency also predicted occupancy, others were markedly different, with competition- and dispersal-related traits becoming more important for occupancy. Which traits were important differed for frequency and occupancy and also differed depending on patch management, wetness, and isolation.

Conclusions

Plant traits can provide insight into plant distribution in fragmented landscapes and can reveal specific abiotic, biotic, and dispersal processes affecting species occurrence in a patch type. However, the importance of individual traits depends on the type of suitable patches available within the landscape.

     

Estimating density of a territorial species in a dynamic landscape

Context

Conservation planning for at-risk species requires understanding of where species are likely to occur, how many individuals are likely to be supported on a given landscape, and the ability to monitor those changes through time.

Objectives

We developed a distribution model for northern spotted owls that incorporates both habitat suitability and probability of territory occupancy while accounting for interspecies competition.

Methods

We developed range-wide habitat suitability maps for two time periods (1993 and 2012) for northern spotted owls that accounted for regional differences in habitat use and home range size. We used these maps for a long-term demographic monitoring study area to assess habitat change and estimate the number of potential territories based on available habitat for both time periods. We adjusted the number of potential territories using known occupancy rates to estimate owl densities for both time periods. We evaluated our range-wide habitat suitability model using independent survey data.

Results

Our range-wide habitat maps predicted areas suitable for territorial spotted owl presence well. On the demographic study area, the amount of habitat declined 19.7% between 1993 and 2012, while our estimate of the habitat-based carrying capacity declined from 150 to 146 territories. Estimated number of occupied territories declined from 94 to 57.

Conclusions

Conservation and recovery of at-risk species depends on understanding how habitat changes over time in response to factors such as wildfire, climate change, biological invasions, and interspecies competition, and how these changes influence species distribution. We demonstrate a model-based approach that provides an effective planning tool.

     

Interactive effects of landscape-wide intensity of farming practices and landscape complexity on wild bee diversity

Context

The local intensity of farming practices is considered as an important driver of biodiversity in agricultural landscapes and its effect on biodiversity has been shown to interact with landscape complexity. But the influence of landscape-wide intensity of farming practices on biodiversity and its combined effect with landscape complexity have been little explored.

Objective

In this study, we tested the interactive effect of the landscape-wide intensity of farming practices and landscape complexity on the local species richness and abundance of farmland wild bee communities.

Methods

We captured wild bees in 96 crop fields and explored the effect of landscape-wide intensity of various farming practices along a gradient of landscape complexity (proportion of semi-natural habitats).

Results

We found that species richness and abundance of wild bees were more positively influenced by landscape complexity in highly insecticide-sprayed landscapes than in less intensively managed landscapes. In contrast, we found that the positive effect of landscape complexity on bee species richness only occurred in landscapes with low nitrogen inputs.

Conclusions

Our study demonstrates the interactive effects of landscape-wide farming intensity and landscape complexity in shaping the diversity of farmland wild bee communities. We conclude that the management of farming intensity at the landscape-scale could mitigate the effects of habitat loss on wild bee decline and would help to maintain pollination services in agricultural landscapes.

     

Consequences of a large-scale fragmentation experiment for Neotropical bats: disentangling the relative importance of local and landscape-scale effects

Context

Habitat loss, fragmentation and degradation are widespread drivers of biodiversity decline. Understanding how habitat quality interacts with landscape context, and how they jointly affect species in human-modified landscapes, is of great importance for informing conservation and management.

Objectives

We used a whole-ecosystem manipulation experiment in the Brazilian Amazon to investigate the relative roles of local and landscape attributes in affecting bat assemblages at an interior-edge-matrix disturbance gradient.

Methods

We surveyed bats in 39 sites, comprising continuous forest (CF), fragments, forest edges and intervening secondary regrowth. For each site, we assessed vegetation structure (local-scale variable) and, for five focal scales, quantified habitat amount and four landscape configuration metrics.

Results

Smaller fragments, edges and regrowth sites had fewer species and higher levels of dominance than CF. Regardless of the landscape scale analysed, species richness and evenness were mostly related to the amount of forest cover. Vegetation structure and configurational metrics were important predictors of abundance, whereby the magnitude and direction of response to configurational metrics were scale-dependent. Responses were ensemble-specific with local-scale vegetation structure being more important for frugivorous than for gleaning animalivorous bats.

Conclusions

Our study indicates that scale-sensitive measures of landscape structure are needed for a more comprehensive understanding of the effects of fragmentation on tropical biota. Although forest fragments and regrowth habitats can be of conservation significance for tropical bats our results further emphasize that primary forest is of irreplaceable value, underlining that their conservation can only be achieved by the preservation of large expanses of pristine habitat.

     

Scale-specific habitat relationships influence patch occupancy: defining neighborhoods to optimize the effectiveness of landscape-scale grassland bird conservation

Context

Beyond the recognized importance of protecting large areas of contiguous habitat, conservation efforts for many species are complicated by the fact that patch suitability may also be affected by characteristics of the landscape within which the patch is located. Currently, little is known about the spatial scales at which species respond to different aspects of the landscape surrounding an occupied patch.

Objectives

Using grassland bird point count data, we describe an approach to evaluating scale-specific effects of landscape composition on patch occupancy.

Methods

We used data from 793 point count surveys conducted in idle and grazed grasslands across Wisconsin, USA from 2012 to 2014 to evaluate scale-dependencies in the response of grassland birds to landscape composition. Patch occupancy models were used to evaluate the relationship between occupancy and landscape composition at scales from 100 to 3000 m.

Results

Bobolink (Dolichonyx oryzivorus) exhibited a pattern indicating selection for grassland habitats in the surrounding landscape at all spatial scales while selecting against other habitats. Eastern Meadowlark (Sturnella magna) displayed evidence of scale sensitivity for all habitat types. Grasshopper Sparrow (Ammodramus savannarum) showed a strong positive response to pasture and idle grass at all scales and negatively to cropland at large scales. Unlike other species, patch occupancy by Henslow’s Sparrow (A. henslowii) was primarily influenced by patch area.

Conclusions

Our results suggest that both working grasslands (pasture) and idle conservation grasslands can play an important role in grassland bird conservation but also highlight the importance of considering species-specific patch and landscape characteristics for effective conservation.

     

Island biogeography theory outweighs habitat amount hypothesis in predicting plant species richness in small grassland remnants

Context

The habitat amount hypothesis has rarely been tested on plant communities. It remains unclear how habitat amount affect species richness in habitat fragments compared to island effects such as isolation and patch size.

Objectives

How do patch size and spatial distribution compared to habitat amount predict plant species richness and grassland specialist plant species in small grassland remnants? How does sampling area affect the prediction of spatial variables on species richness?

Methods

We recorded plant species density and richness on 131 midfield islets (small remnants of semi-natural grassland) situated in 27 landscapes in Sweden. Further, we tested how habitat amount, compared to focal patch size and distance to nearest neighbor predicted species density and richness of plants and of grassland specialists.

Results

A total of 381 plant species were recorded (including 85 grassland specialist species). A combination of patch size and isolation was better in predicting both density and richness of species compared to habitat amount. Almost 45% of species richness and 23% of specialist species were explained by island biogeography parameters compared to 19 and 11% by the amount of habitat. A scaled sampling method increased the explanation level of island biogeography parameters and habitat amount.

Conclusions

Habitat amount as a concept is not as good as island biogeography to predict species richness in small habitats. Priority in landscape planning should be on larger patches rather than several small, even if they are close together. We recommend a sampling area scaled to patch size in small habitats.

     

Integrating a generic focal species,metapopulation capacity,and connectivity to identify opportunities to link fragmented habitat

Context

A challenge devising revegetation strategies in fragmented landscapes is conserving for the widest spectrum of biodiversity. Habitat network reconstruction should improve landscape capacity to maintain species populations. However, the location of revegetation often fails to account for species occurrence and dispersal processes operating across spatial scales.

Objectives

Our objective was to integrate metapopulation theory with estimates of landscape capacity and dispersal pathways to highlight connectivity gaps. Maintenance of populations could thereby be facilitated through reconnecting habitat networks across regional and broader scales, with assumed benefit for the dispersal needs of less sensitive species.

Methods

Predicted occupancy and metapopulation capacity were calculated for a generic focal species derived from fragmentation-sensitive woodland birds, mammals and reptiles. A metapopulation connectivity analysis predicted regional dispersal links to identify likely routes through which individuals may move to contribute to the viability of the population. We used the revegetation programmes of the Brigalow–Nandewar Biolinks project, eastern New South Wales, Australia, to demonstrate our approach.

Results

Landscape capacity of the current landscape varied across the region. Low-value links between populations provided greatest opportunities for revegetation and improved landscape capacity. Where regional connectivity did not indicate a pathway between populations, broader scale connectivity provided guidance for revegetation.

Conclusions

The metapopulation-based model, coupled with a habitat dispersal network analysis, provided a platform to inform revegetation locations and better support biodiversity. Our approach has application for directing on-ground action to support viable populations, assess the impact of revegetation schemes or monitor the progress of staged implementations.

     

Synthetic Aperture Radar (SAR) images improve habitat suitability models

Context

The ability to detect ecological networks in landscapes is of utmost importance for managing biodiversity and planning corridors.

Objectives

The objective of this study was to evaluate the information provided by a synthetic aperture radar (SAR) image for landscape connectivity modeling compared to aerial photographs (APs).

Methods

We present a novel method that integrates habitat suitability derived from remote sensing imagery into a connectivity model to explain species abundance. More precisely, we compared how two resistance maps constructed using landscape and/or local metrics derived from AP or SAR imagery yield different connectivity values (based on graph theory), considering hedgerow networks and forest carabid beetle species as a model.

Results

We found that resistance maps using landscape and local metrics derived from SAR imagery improve landscape connectivity measures. The SAR model is the most informative, explaining 58% of the variance in forest carabid beetle abundance. This model calculates resistance values associated with homogeneous patches within hedgerows according to their suitability (canopy cover density and landscape grain) for the model species.

Conclusions

Our approach combines two important methods in landscape ecology: the construction of resistance maps and the use of buffers around sampling points to determine the importance of landscape factors. This study was carried out through an interdisciplinary approach involving remote sensing scientists and landscape ecologists. This study is a step forward in developing landscape metrics from satellites to monitor biodiversity.

     

Bat richness and activity in heterogeneous landscapes: guild-specific and scale-dependent?

Context

The conversion of natural environments into agricultural land has profound effects on the composition of the landscape, often resulting in a mosaic of human-altered and natural habitats. The response to these changes may however vary among organisms. Bats are highly vagile, and their requirements often imply the use of distinct habitats, which they select responding to both landscape and local features.

Objectives

We aimed to identify which features influence bat richness and activity within Baixo Vouga Lagunar, a heterogeneous landscape located on the Central-North Portuguese coast, and to investigate if that influence varies across a gradient of focal scales.

Methods

We sampled bats acoustically, while simultaneously sampling insects with light traps. We assessed the relationships between species richness, bat activity, and activity of eco-morphological guilds with landscape and local features, across four scales.

Results

Our results revealed both scale- and guild-dependent responses of bats to landscape and local features. At broader scales we found positive associations between open-space foraging bats and habitat heterogeneity and between edge-space foraging bats and greater edge lengths. Woodland cover and water availability at an intermediate scale and weather conditions and insect abundance at a local scale were the factors that mostly influenced the response variables.

Conclusions

Globally, our results suggest that bats are sensitive to local resource availability and distribution, while simultaneously reacting to landscape features acting at coarser scales. Finally, our results suggest that the responses given by bats are guild-dependent, and some habitats act as keystone structures for bats within this mosaic.

     

Metapopulation modelling of long-term urban habitat-loss scenarios

Context

Increasing human populations in urban areas pose a threat to species’ persistence through habitat loss and fragmentation. It is therefore essential that we develop methods to investigate critical habitat loss thresholds and least detrimental landscape configurations.

Objectives

We develop a framework to assess how the pattern of habitat loss impacts the ecological and social characteristics of a landscape and how this varies depending on the species and criteria by which it is judged.

Methods

We use a scenario-based approach to test six propositions in which habitat is lost preferentially based on patch characteristics. We use eight bird and two amphibian species as indicator species. To compare scenarios, we present a method combining the output from a metapopulation model with measures of social impacts of land-cover change in a multiple criteria decision analysis. We also determine whether a habitat loss threshold exists, below which small loss of habitat can lead to large loss of species’ occupancy.

Results

We found that, of the scenarios presented, preferentially losing common habitats and smaller patches was least detrimental for both ecological and social factors. Threshold effects were found for all but the generalist bird species.

Conclusions

We have outlined a workflow which allows for transparent, repeatable comparison between landscapes. This workflow can be used to compare urban landscape plans, or to develop general understanding of the impacts of different forms of habitat loss. Reassuringly, the recommendations based on the scenarios presented are in keeping with received conservation wisdom: to prioritise larger and/or rarer patches.

     

Distribution of invasive plants in urban environment is strongly spatially structured

Context

Urban environments create a wide range of habitats that harbour a great diversity of plant species, many of which are of alien origin. For future urban planning and management of the green areas within the city, understanding of the spatial distribution of invasive alien species is of great importance.

Objectives

Our main aim was to assess how availability of different ecosystem types within a city area, as well as several parameters describing urban structure interact in determining the cover and identity of invasive alien species.

Methods

We studied the distribution of chosen invasive plant species in a mid-sized city in the Czech Republic, central Europe, on a gradient of equal sized cells from the city centre to its outskirts.

Results

A great amount of variation was explained by spatial predictors but not shared with any measured variables. The species cover of invasive species decreased with increasing proportion of urban greenery and distance from the city centre, but increased with habitat richness; road margins, ruderal sites, and railway sites were richest in invasive species. In contrast, the total number of invasive species in cells significantly decreased with increasing distance from the city centre, but increased with habitat richness.

Conclusions

Our results suggest that different invasive species prefer habitats in the vicinity of the city centre and at its periphery and the spatial structure and habitat quality of the urban landscape needs to be taken into account, in efforts to manage alien plant species invasions in urban environments.

     

Assessing wild bees in perennial bioenergy landscapes: effects of bioenergy crop composition,landscape configuration,and bioenergy crop area

Context

Wild bee populations are currently under threat, which has led to recent efforts to increase pollinator habitat in North America. Simultaneously, U.S. federal energy policies are beginning to encourage perennial bioenergy cropping (PBC) systems, which have the potential to support native bees.

Objectives

Our objective was to explore the potentially interactive effects of crop composition, total PBC area, and PBC patches in different landscape configurations.

Methods

Using a spatially-explicit modeling approach, the Lonsdorf model, we simulated the impacts of three perennial bioenergy crops (PBC: willow, switchgrass, and prairie), three scenarios with different total PBC area (11.7, 23.5 and 28.8% of agricultural land converted to PBC) and two types of landscape configurations (PBC in clustered landscape patterns that represent realistic future configurations or in dispersed neutral landscape models) on a nest abundance index in an Illinois landscape.

Results

Our modeling results suggest that crop composition and PBC area are particularly important for bee nest abundance, whereas landscape configuration is associated with bee nest abundance at the local scale but less so at the regional scale.

Conclusions

Strategies to enhance wild bee habitat should therefore emphasize the crop composition and amount of PBC.

     

Stop using dichotomous terms to reference observations of scale-dependent habitat selection

Context

Many studies dichotomize habitat selection into “local” or “landscape” effects, with little explanation regarding what each represents. Ambiguous use of these terms across studies may confound observations of selection operating across different spatiotemporal resolutions and impede synthesis of scale-dependent habitat selection.

Objective

To examine the consistency of use of the terms “local” and “landscape” level habitat selection and evaluate potential implications for conservation science.

Methods

We reviewed 136 multi-level studies observing local and/or landscape effects. From each study, we identified which of Johnson’s (Ecology 61:65–71, 1980) selection orders was observed at the local and landscape scale. We assessed consistency in the selection order observed at each scale.

Results

At the local scale, 54% of studies observed 3rd order selection and 38% of studies observed 4th order selection. At the landscape scale, 64% of studies observed 2nd order selection, while only 8% of studies observed 1st order selection. Overlap also occurred; 2nd order selection was observed at the local scale in 8% of studies, while 3rd and 4th order selection was observed at the landscape scale in 24 and 3% of studies, respectively.

Conclusions

Selection orders observed at local and landscape scales are inconsistent across studies, making comparisons and synthesis difficult. This inconsistency makes the terms “local” and “landscape” meaningless, and may lead to misidentification of limiting factor(s) most important for conservation efforts. Dichotomous terms should no longer be used in reference to levels of selection, but in reference to specific explanatory variables whose characteristic scale(s) best fit the appropriate definition.

     

Can waterbirds with different movement,dietary and foraging functional traits occupy similar ecological niches?

Context

Competitive interactions potentially play an important role in structuring bird communities. It is unclear how differences in functional traits influence the niche dimensions of highly mobile waterbird species, particularly when they co-exist in spatiotemporally heterogeneous communities.

Objectives

We investigated the inter-relationships between waterbird trait groupings (movement, dietary and foraging habitat) and environmental variable groupings (rainfall, land cover, vegetation structure and water quality). Specifically, we tested whether the scale of environmental variables filtered movement traits and whether these traits operated in conjunction with dietary and foraging habitat traits to form distinct ecological niches in waterbirds.

Methods

We conducted waterbird and environmental variable surveys in 60 sites, sampled seven times each at bimonthly intervals, in KwaZulu-Natal, South Africa. Trait-environment relationships were tested using a combination of RLQ and fourth-corner analyses.

Results

Several significant trait-environment relationships emerged in bivariate correlations and multivariate ordination space. Movement traits correlated with the scale of environmental variables; migrant and nomadic species responded to broad scale environmental variables. Vegetation structure and land cover were particularly important in explaining the abundance of species foraging in emergent vegetation. Three groups emerged along a gradient in multivariate ordination space providing evidence for ecological niche separation of waterbirds with different movement traits.

Conclusions

Our findings suggest that the scale of landscape resources can act as a filter of movement traits, and that in conjunction with dietary and foraging traits, waterbirds with different movement traits occupy distinct ecological niches.

     

A multi-species approach for assessing the impact of land-cover changes on landscape connectivity

Context

Land-cover changes (LCCs) could impact wildlife populations through gains or losses of natural habitats and changes in the landscape mosaic. To assess such impacts, we need to focus on landscape connectivity from a diachronic perspective.

Objectives

We propose a method for assessing the impact of LCCs on landscape connectivity through a multi-species approach based on graph theory. To do this, we combine two approaches devised to spatialize the variation of multi-species connectivity and to quantify the importance of types of LCCs for single-species connectivity by highlighting the possible contradictory effects.

Methods

We begin with a list of landscape species and create virtual species with similar ecological requirements. We model the ecological network of these virtual species at two dates and compute the variation of a local and global connectivity metric to assess the impacts of the LCCs on their dispersal capacities.

Results

The spatial variation of multi-species connectivity showed that local impacts range from ?6.4% to +3.2%. The assessment of the impacts of types of LCCs showed a variation in global connectivity ranging from ?45.1% for open-area reptiles to +170.2% for natural open-area birds with low-dispersion capacities.

Conclusions

This generic approach can be reproduced in a large variety of spatial contexts by adapting the selection of the initial species. The proposed method could inform and guide conservation actions and landscape management strategies so as to enhance or maintain connectivity for species at a landscape scale.

     

Conservation planning with spatially explicit models: a case for horseshoe bats in complex mountain landscapes

Context

Context Bats are considered as an ecological indicator of habitat quality due to their sensitivity to human-induced ecosystem changes. Hence, we will focus the study on two indicator species of bats as a proxy to evaluate structure and composition of the landscape to analyze anthropic pressures driving changes in patterns.

Objectives

This study develops a spatially-explicit model to highlight key habitat nodes and corridors which are integral for maintaining functional landscape connectivity for bat movement. We focus on a complex mountain landscape and two bat species: greater (Rhinolophus ferrumequinum) and lesser (Rhinolophus hipposideros) horseshoe bats which are known to be sensitive to landscape composition and configuration.

Methods

Species distribution models are used to delineate high-quality foraging habitat for each species using opportunistic ultrasonic bat data. We then performed connectivity analysis combining (modelled) suitable foraging habitat and (known) roost sites. We use graph-theory and the deviation in the probability of connectivity to quantify resilience of the landscape connectivity to perturbations.

Results

Both species were confined to lowlands (<1000 m elevation) and avoided areas with high road densities. Greater horseshoe bats were more generalist than lesser horseshoe bats which tended to be associated with broadleaved and mixed forests.

Conclusions

The spatially-explicit models obtained were proven crucial for prioritizing foraging habitats, roost sites and key corridors for conservation. Hence, our results are being used by key stakeholders to help integrate conservation measures into forest management and conservation planning at the regional level. The approach used can be integrated into conservation initiatives elsewhere.

     

Employing landscape metrics in an open population model to estimate demographic parameters of a grassland bird

Context

The importance of landscape context is increasingly recognized when studying relationships between populations. Recent advances in open population modeling allow the employment of landscape metrics to estimate demographic parameters underlying population variation through time and space.

Objectives

Our primary objectives were to (1) describe the influence of landscape metrics on demographic parameters in the grasshopper sparrow (Ammodramus savannarum) and (2) quantify the contributions of these demographic parameters in influencing variation in territory counts through time. We anticipated results would allow us to make recommendations for prioritizing site conservation for this grassland-obligate species of regional conservation concern.

Methods

We employed territory counts spanning 13 years from Massachusetts, USA in open population models to estimate the effects of landscape metrics, territory density, and site quality on three demographic parameters.

Results

The best model estimated highest initial numbers of territories in larger, more distant sites. Overall growth rates <1 were estimated during 1993–2005, while growth rates >1 were estimated in larger sites with a higher habitat quality index and low to medium relative density. Highest rates of annual immigration were estimated for larger sites. Growth rate explained the greatest proportion of variation in territory counts through time.

Conclusions

Open population models allowed us to identify the effects of landscape context on multiple grasshopper sparrow demographic parameters. We encourage further application of these and related models to grassland birds. Beyond maintaining grasslands in the region, we recommend the conservation of large, distant, and previously occupied sites to benefit regional populations.