Nesting Success of a Songbird in a Complex Floodplain Forest Landscape in llinois, USA: Local Fragmentation vs. Vegetation Structure

Measuring edge effects in complex landscapes is often confounded by the presence of different kinds of natural and anthropogenic edges, each of which may act differently on organisms inhabiting habitat patches. In such landscapes, proportions of different habitats surrounding nests within patches often vary and may affect nesting success independently of distance to edges. We developed methods to measure and study the effects of multiple edges and varying habitat composition around nests on the breeding success of the Acadian flycatcher (Empidonax virescens), an understory, open-cup nesting songbird. The Kaskaskia River in Southwestern Illinois was our study area and consists of wide (>1000-m) floodplain corridors embedded in an agricultural matrix with a variety of natural (wide rivers, backwater swamps, and oxbow lakes) and anthropogenic (internal openings, and agricultural) habitats. We also measured vegetation structure around each nest. Nest survival increased with increasing nest concealment, and probabilities of brood parasitism increased with increasing distances from anthropogenic and natural water-related openings surrounding nests. The magnitude of these effects was small, probably because the landscape is saturated with nest predators and brood parasites. These results illustrate the importance of considering both larger landscape context and details of natural and anthropogenic disturbances when studying the effects of habitat fragmentation on wildlife.     

Modelling the Effects of Dispersal and Landscape Configuration on Population Distribution and Viability in Fragmented Habitat

Landscape configuration and dispersal characteristics are major determinants of population distribution and persistence in fragmented habitat. An individual-based spatially explicit population model was developed to investigate these factors using the distribution of nuthatches in an area of eastern England as an example. The effects of immigration and increasing the area of breeding quality habitat were explored. Predictions were compared with observed population sizes in the study area. Our model combined a nuthatch population simulator based on individual behaviour with a grid-based representation of the landscape; nuthatch life cycle and immigration parameters were user selectable. A novel aspect of the model is user-selection of habitat perceptual range. Using a realistic set of parameters, the number of breeding pairs predicted by the model matched observed numbers. According to model simulations, the main cause of nuthatch scarcity in the study area was the inability of patches to support viable populations without immigration from elsewhere. Modelled habitat management, which increased breeding quality habitat in existing woods, lowered the threshold above which the study area population became self-sustaining. The existence of a large core habitat area was critical in producing a self-sustaining population in this landscape, the same area in dispersed small woods failed to sustain populations.     

Sacrificing patches for linear habitat elements enhances metapopulation performance of woodland birds in fragmented landscapes

It is generally assumed that large patches of natural habitat are better for the survival of species than the same amount of habitat in smaller fragments or linear elements like hedges and tree rows. We use a spatially explicit individual-based model of a woodland bird to explore this hypothesis. We specifically ask whether mixtures of large, small and linear habitat elements are better for population performance than landscapes that consist of only large elements. With equal carrying capacity, metapopulations perform equally or better in heterogeneous landscape types that are a mix of linear, large and small habitat elements. We call this increased metapopulation performance of large and small elements “synergy”. These mixed conditions are superior because the small linear elements facilitate dispersal while patches secure the population in the long run because they have a lower extinction risk. The linear elements are able to catch and guide dispersing animals which results in higher connectivity between patches leading to higher metapopulation survival. Our results suggest that landscape designers should not always seek to conserve and create larger units but might better strive for more variable landscapes with mixtures of patch sizes and shapes. This is especially important when smaller units play a key role in connecting patches and dispersal through the matrix is poor.     

Effects of Habitat Suitability and Landscape Patterns on Tick (Acarina) Metapopulation Processes

Tick density and population dynamics are important factors in the ecological processes involved in pathogen circulation in a habitat. These characteristics of tick populations are closely linked to habitat suitability, which reflects the limiting ecological factors and landscape features affecting tick populations; however, little work has been done on the regional assessment of habitat suitability. In this study, a regional model for the distribution and abundance of the tick Ixodes ricinus in central Spain is developed. An occurrence and an abundance model were constructed; climate and vegetation variables were found to be the main predictors of both occurrence and density in a relatively homogeneous matrix of habitat patches, whereas topographical variables were found to have small contributions and were therefore discarded. The residuals of the abundance model showed good correlation with the isolation of each patch. The predictive power of the abundance model was greatly enhanced by inclusion of the traversability (a measure of the permeability of each patch to the propagules of the metapopulation) and recruitment (an index of the relative importance of each patch to the traffic through the entire habitat network). The removal from the landscape of the patches whose recruitment values were in the top 10% has a critical effect on tick density, an effect not observed when patches are removed at random. These results indicate that permanent tick populations can be sustained only in landscapes containing a minimum network of viable sites. Graph theory and measurements of patch isolation should prove to be important elements in the forecasting of tick abundance and the management of the features underlying the landscape ecology of tick populations and pathogen circulation in the field.     

The influence of urbanisation on diversity and trait composition of birds

We analysed the effect of the urban matrix, the urban space surrounding distinct habitat patches, on bird communities. In doing so we assessed the impact of urbanisation beyond the effect of habitat loss. We used a set of 54 wasteland sites of early successional stages that were scattered over the entire urban area of Berlin, Germany. Sites were similar to each other in habitat structure but differed in their surroundings, the urban matrix. Thus, our study design allows to investigate associations between birds and the urban matrix. Our measures for urbanisation are human population density and degree of sealing within 50 to 2,000 m buffer zones surrounding each wasteland site. Along the urbanisation gradients we calculated three measures of bird communities: alpha diversity, beta diversity, and trait profile of the entire bird community regarding food, life-history, and behavioural traits. Alpha diversity did not change significantly along the gradients of urbanisation. However, beta diversity increased along the urbanisation gradients with urbanisation at the local scale (50 m) but decreased at the landscape scale (200 and 2,000 m). Fourth-corner analysis of relationships between urbanisation and species traits showed trait shifts: adult survival rate increased with human population density and densities of birds that are more often reported to show innovative behaviour increased with both human population density and degree of sealing. We conclude that the influence of the urban matrix contributes to the homogenisation of the avifauna by filtering certain species traits and promoting others.     

The costs of multiple patch use by birds

Birds living in fragmented habitat may occupy territories comprising more than one patch. This paper uses a theoretical model to investigate the costs (in terms of time and energy) of crossing gaps between patches for birds feeding young in the nest, using the great tit (Parus major) as an example. When the proportion of foraging trips involving gap-crossing was small (25%), gaps of about 300–550 m (depending on body mass and flight speed) could be crossed without exceeding likely maximum sustainable daily energy expenditure (DEE_max). However, a penalty of time lost in crossing gaps of about one hour was incurred. For more gap-crossing (due to larger brood size and/or a greater proportion of gap-crossing trips), distances that could be crossed decreased rapidly to about 50–100 m and time lost increased to more than six hours. Crossing gaps at maximum range speed, rather than at the slower minimum power speed, reduced flight times by 42% and slightly reduced overall daily energy expenditure because the higher flight costs per minute were more than off-set by the shorter flight times. Smaller body mass (17 g versus 19 g) was advantageous for gap-crossing, the distances which could be crossed without exceeding DEE_max being almost doubled for the smaller mass. The influence of changes in wing morphology, fat load and prey load size on the energetics of gap-crossing were also considered. Although the model was constructed for a woodland bird, problems of time and energy expenditure associated with gap-crossing will affect many species which exploit patchy resources, especially when the spacing of the patches increases, for example due to habitat loss and modification. In landscapes where semi-natural habitat is highly fragmented and most surviving patches are small (e.g., many farming landscapes) the costs of multiple patch use may represent another mechanism by which habitat fragmentation reduces the reproductive potential of the inhabitants of habitat patches which are of acceptable or even good quality, but are small.     

The Effects of Land-use and Landscape Structure on Barn Owl (<Emphasis Type="Italic">Tyto alba</Emphasis>) Breeding Success in Southern England,U.K.

To aid effective conservation and management there is a need to understand the effect of landscape on species ecology. The aim of this research was to assess the effect of landscape parameters on breeding success of barn owls throughout the Rother and Arun River catchments, Sussex, UK. We used a Geographic Information System to describe the habitat mosaic and landscape structure within an estimated home range area of 3 km² around 85 artificial nest box sites. Results showed that land cover was less heterogeneous at successful sites, with home ranges dominated by a few habitat types of regular patch shapes. Unsuccessful nesting sites had significantly more improved grassland, suburban land and wetlands than successful sites. Cluster analysis and Principle Components Analysis was used to assess the similarity of the habitat mosaic within these areas and pellet analysis was undertaken to assess barn owl diet and prey availability. Ten prey species were recovered from pellets, field vole (Microtus agrestis), common shrews (Sorex araneus) and house mice (Mus musculus) making up nearly 90% of recoveries. However box sites varied in relative proportions of small mammal, and hence prey availability. Results indicated that land use and landscape structure can affect breeding success in barn owls. Higher levels of poor quality small mammal habitat were associated with unsuccessful sites. However, at a landscape scale, the habitat mosaic across the study area lacked variation, limiting analysis and clear correlations between habitat type and positive breeding success, suggesting that a finer scale was needed in future studies utilising this approach.     

Analyzing the effect of stepping stones on target patch colonisation in structured landscapes for Eurasian lynx

With habitat loss and fragmentation having become two of the major threats to the viability of species, the question of how to manage landscapes for species conservation has attracted much attention. In this context, the planning of stepping stones has been proposed to increase connectivity in fragmented landscapes. We present a simulation study with a neutral landscape approach to assess the effects of stepping stones on colonization success. To that end, we used a spatially explicit, calibrated population model of the European lynx (Lynx lynx) coupled with structured landscapes, in which we could control the landscape parameters of dispersal habitat coverage and contagion, as well as the number and size of stepping stones available for breeding. In general, we found that colonization success increased with increasing habitat coverage but decreased with increasing habitat contagion, while the introduction of stepping stones had significant effects in critical situations. Especially at low to medium dispersal habitat coverage and high disperser mortality, stepping stones had a positive effect on colonization success when they were large enough to produce new dispersers, but negative effects when they were small and located in a way that dispersers would be distracted from more suitable breeding habitat patches. The latter clearly constituted a shading effect and argues for a thorough consideration of the trade-offs related to stepping stone size and location when implementing stepping stones as a conservation measure, especially when the number of individuals of conservation concern is low.     

Predator activity and predation on songbird nests on forest-field edges in east-central Illinois

We measured the activity of mammalian predators, numbers of singing male songbirds, and predation rates on nests of songbirds (152 natural, open-cup nests and 380 artificial nests) on 38 250 m transects located along various types of forest-field edges in a wildlife management area in east-central Illinois. We then related these variables to each other and to measures of the vegetative structure of our transects that we anticipated might influence predator activity or predation rates on nests of birds characteristic of edge and shrubland habitats. Mammalian predators, particularly raccoons (Procyon lotor), were abundant in the wildlife area and present on all transects surveyed. We did not find significant relationships among the variables we measured. Rather, rates of nest predation were consistently high (>70%) and generally evenly distributed around our study site. Medium-sized, generalist mammalian predators in the midwestern United States reach their highest population densities in fragmented landscapes with abundant edge habitat, particularly agricultural edges. Areas of natural habitat in these landscapes dominated by agriculture may concentrate predators and act as ecological traps for nesting birds because they attract high densities of breeding birds that are subjected to high rates of nest predation.     

Landscape-scale Edge Effect in Predation Risk in Forest-farmland Mosaics of Central Europe

Avian nest predation is known to increase with the degree of forest fragmentation. A common explanation is that farmland allows for high densities of generalist predators, and predators penetrating into the forest cause higher nest losses at forest-farmland edges than in forest interiors. In contrast to numerous patch-level studies of forest edge effects conducted earlier, we broadened the spatial extent to the landscape. We tested the hypothesis of increased predation near farmland over distances of >4 km from forest–farmland edges into forest interiors in five mountain ranges in Germany, using artificial ground nests. We considered two landscape settings: (1) Transitions between a forest matrix and a farmland matrix, and (2) farmland patches within a forest matrix. Nest losses were not significantly higher in vicinity to a farmland matrix, but proximity to a pasture within the forest matrix strongly increased predation risk. We speculate that these differences resulted from landscape geometry. Farmland patches and matrix alike are highly attractive to generalist predators, and are regularly visited by red foxes from the forest. Predators that traverse the forest and take prey along the way, will cause a concentration of predation risk towards a patch (pasture), but not towards an adjacent matrix (farming lowlands), of feeding habitat. Contrary to previous evidence that edge effects in nest predation level off after 50 m, nest fate was related to distance to pastures across the entire study extent of 4.1 km. Our results suggest that landscape context and predator mobility may greatly affect spatial predation patterns.     

Population dynamics and habitat connectivity affecting the spatial spread of populations – a simulation study

In this paper we show how the spatialconfiguration of habitat quality affects the spatial spread of apopulation in a heterogeneous environment. Our main result is thatfor species with limited dispersal ability and a landscape withisolated habitats, stepping stone patches of habitat greatlyincrease the ability of species to disperse. Our results showthat increasing reproductive rate first enables and thenaccelerates spatial spread, whereas increasing the connectivity has aremarkable effect only in case of low reproductive rates. Theimportance of landscape structure varied according to thedemographic characteristics of the population. To show this wepresent a spatially explicit habitat model taking into accountpopulation dynamics and habitat connectivity. The population dynamicsare based on a matrix projection model and are calculated on eachcell of a regular lattice. The parameters of the Leslie matrix dependon habitat suitability as well as density. Dispersal between adjacentcells takes place either unrestricted or with higher probability inthe direction of a higher habitat quality (restricted dispersal).Connectivity is maintained by corridors and stepping stones ofoptimal habitat quality in our fragmented model landscape containinga mosaic of different habitat suitabilities. The cellular automatonmodel serves as a basis for investigating different combinations ofparameter values and spatial arrangements of cells with high and lowquality.This revised version was published online in May 2005 with corrections to the Cover Date.     

How to define a patch: a spatial model for hierarchically delineating organism-specific habitat patches

Landscape analysis and delineation of habitat patches should take into account organism-specific behavioral and perceptual responses to landscape structure because different organisms perceive and respond to landscape features over different ranges of spatial scales. The commonly used methods for delineating habitat based on rules of contiguity do not account for organism-specific responses to landscape patch structure and have undesirable properties, such as being dependent on the scale of base map used for analysis. This paper presents an improved patch delineation algorithm, “PatchMorph,” which can delineate patches across a range of spatial scales based on three organism-specific thresholds: (1) land cover density threshold, (2) habitat gap maximum thickness (gap threshold), and (3) habitat patch minimum thickness (spur threshold). This algorithm was tested on an “idealized” landscape with landscape gaps and spurs of known size, and delineated patches as expected. It was then applied to delineate patches from a neutral random fractal landscape, which showed that as the input gap and spur thickness thresholds were increased, the number of patches decreased from 59 (low thresholds) patches to 1 (high thresholds). The algorithm was then applied to model western yellow-billed cuckoo (Coccyzus americanus occidentalis) nesting habitat patches based on spur and gap thresholds specific to this organism. Both these analyses showed that fewer patches were delineated by PatchMorph than by rules of contiguity, and those patches were larger, had smoother edges, and had fewer gaps within the patches. This algorithm has many applications beyond those presented in this paper, including habitat suitability analysis, spatially explicit population modeling, and habitat connectivity analysis.     

Response of mountain plovers to plague-driven dynamics of black-tailed prairie dog colonies

Sylvatic plague is a major factor influencing the dynamics of black-tailed prairie dog (Cynomys ludovicianus) colonies in the western Great Plains. We studied the nesting response of the mountain plover (Charadrius montanus), a grassland bird that nests on prairie dog colonies, to plague-driven dynamics of prairie dog colonies at three sites in the western Great Plains. First, we examined plover nest distribution on colonies that were previously affected by plague, but that had been recovering (expanding) for at least 6 years. Plovers consistently nested in both young (colonized in the past 1–2 years) and old (colonized for 6 or more years) portions of prairie dog colonies in proportion to their availability. Second, we examined changes in plover nest frequency at two sites following plague epizootics, and found that mountain plover nest numbers declined relatively rapidly (≤2 years) on plague-affected colonies. Taken together, our findings indicate that available plover nesting habitat associated with prairie dog colonies closely tracks the area actively occupied by prairie dogs each year. Given the presence of plague throughout most of the mountain plover’s breeding range in the western Great Plains, important factors affecting plover populations likely include landscape features that determine the scale of plague outbreaks, the distance that plovers move in response to changing breeding habitat conditions, and the availability and quality of alternate breeding habitat within the landscape.     

Habitat Selection and Breeding Success in a Forest-nesting Alcid, the Marbled Murrelet, in Two Landscapes with Different Degrees of Forest Fragmentation

We studied habitat selection and breeding success in marked populations of a protected seabird (family Alcidae), the marbled murrelet (Brachyramphus marmoratus), in a relatively intact and a heavily logged old-growth forest landscape in south-western Canada. Murrelets used old-growth fragments either proportionately to their size frequency distribution (intact) or they tended to nest in disproportionately smaller fragments (logged). Multiple regression modelling showed that murrelet distribution could be explained by proximity of nests to landscape features producing biotic and abiotic edge effects. Streams, steeper slopes and lower elevations were selected in both landscapes, probably due to good nesting habitat conditions and easier access to nest sites. In the logged landscape, the murrelets nested closer to recent clearcuts than would be expected. Proximity to the ocean was favoured in the intact area. The models of habitat selection had satisfactory discriminatory ability in both landscapes. Breeding success (probability of nest survival to the middle of the chick rearing period), inferred from nest attendance patterns by radio-tagged parents, was modelled in the logged landscape. Survivorship was greater in areas with recent clearcuts and lower in areas with much regrowth, i.e. it was positively correlated with recent habitat fragmentation. We conclude that marbled murrelets can successfully breed in old-growth forests fragmented by logging.     

Joint effects of habitat configuration and temporal stochasticity on population dynamics

Habitat configuration and temporal stochasticity in the environment are recognized as important drivers of population structure, yet few studies have examined the combined influence of these factors. We developed a spatially explicit simulation model to investigate how stochasticity in survival and reproduction influenced population dynamics on landscapes that differed in habitat configuration. Landscapes ranged from completely contiguous to highly fragmented, and simulated populations varied in mean survival probability (0.2, 0.4, 0.8) and dispersal capacity (1, 3, or 5 cells). Overall, habitat configuration had a large effect on populations, accounting for >80% of the variation in population size when mean survival and dispersal capacity were held constant. Stochasticity in survival and reproduction were much less influential, accounting for <1–14% of the variation in population size, but exacerbated the negative effects of habitat fragmentation by increasing the number of local extinctions in isolated patches. Stochasticity interacted strongly with both mean survival probability and habitat configuration. For example, survival stochasticity reduced population size when survival probability was high and habitat was fragmented, but had little effect on population size under other conditions. Reproductive stochasticity reduced population size irrespective of mean survival and habitat configuration, but had the largest effect when survival probability was intermediate and habitat was well connected. Stochasticity also enhanced the variability of population size in most cases. Contrary to expectations, increasing dispersal capacity did not increase population persistence, because the probability of finding suitable habitat within the dispersal neighborhood declined more for the same level of dispersal capacity when fragmentation was high compared to when it was low. These findings suggest that greater environmental variability, as might arise due to climate change, is likely to compound population losses due to habitat fragmentation and may directly reduce population size if reproductive output is compromised. It may also increase variability in population size.     

Large termitaria act as refugia for tall trees,deadwood and cavity-using birds in a miombo woodland

Landscape heterogeneity can play an important role in providing refugia and sustaining biodiversity in disturbed landscapes. Large Macrotermes (Isoptera) termite mounds in miombo woodlands form nutrient rich islands that sustain a different suite of woody plant species relative to the woodland matrix. We investigated the role of termitaria in providing habitat for cavity-using birds in miombo woodlands that had been greatly impacted by elephants and fire, by comparing the availability of habitat favored by cavity-using birds (tall trees, trees with deadwood, and cavities) on and off mounds, and then testing its effect on species richness and abundance of cavity-using birds. We surveyed 48 termitaria paired with 48 woodland matrix sites in the breeding season; and 54 matrix-termitarium pairs in the non-breeding season in Chizarira National Park, Zimbabwe. Generalized linear mixed-effects models showed that termitaria harboured significantly higher densities (ha⁻¹) of habitat components considered important for cavity nesting birds. Density of trees >6 m in height and incidence of trees with deadwood was nearly 10 times greater on mounds than in the matrix, and the density of cavities was nine times higher on mounds compared to the matrix. A model selection procedure showed that termitaria provided refugia for cavity-using birds and contributed to the resilience of bird communities through high on-mound densities of trees with deadwood. Large termitaria thus appear to play an important role in maintaining functionally important components of the avifauna in heavily impacted Miombo woodlands.     

Disentangling habitat and social drivers of nesting patterns in songbirds

Nest locations of breeding birds are often spatially clustered. This tendency to nest together has generally been related to a patchy distribution of nesting habitat in landscape studies, but behavioral studies of species with clustered breeding patterns draw attention to the importance of social and biotic factors. Indeed, it is becoming increasingly apparent that the breeding system of many territorial, migrant birds may be semi-colonial. The reasons for, and extent of, spatial clustering in their breeding systems are not well understood. Our goal was to tease apart the influence of habitat availability and social drivers of clustered breeding in a neotropical migrant species, the hooded warbler (Wilsonia citrina). To test alternative hypotheses related to clustered habitat or conspecific attraction, we combined a habitat classification based on remote sensing with point pattern analysis of nesting sites. Nest locations (n = 150, 1999–2004), collected in a 1213 ha forested area of Southern Ontario (Canada), were analyzed at multiple spatial scales. Ripley’s K and pair-correlation functions g (uni- and bivariate) were used to test whether nests were clustered merely because potential nesting habitat was also clustered, or whether nests were additionally clustered with respect to conspecifics. Nest locations tended to be significantly clustered at intermediate distances (particularly between 240 and 420 m). Nests were randomly distributed within available habitat at larger distance scales, up to 1500 m. A reasonable hypothesis to explain the detected additional clustering, and one that is consistent with the results of several behavioral studies, is that females pack their nests more tightly than the available habitat requires to be situated closer to their neighbors’ mates. Linking spatially explicit, point pattern analysis with strong inference based on Monte Carlo tests may bring us closer to understanding the generality and reasons behind conspecific attraction at different spatial scales. F. Csillag—deceased.     

Integrating single-species management and landscape conservation using regional habitat occurrence models: the northern goshawk in the Southwest,USA

Conservation planners and land managers are often confronted with scale-associated challenges when assessing the relationship between land management objectives and species conservation. Conservation of individual species typically involves site-level analyses of habitat, whereas land management focuses on larger spatial extents. New models are needed to more explicitly integrate species-specific conservation with landscape or regional scales. We address this challenge with an example using the northern goshawk (Accipiter gentilis), a forest raptor with circumpolar distribution that is the focus of intense debate regarding forest management on public lands in the southwestern USA. To address goshawk-specific habitat conservation across a management area of 22,800-km² in northern Arizona, we focused on the territory scale rather than individual nest sites. We compiled a 17-year database of 895 nest sites to estimate territory locations. We then estimated the likelihood of territory occurrence for the entire management area using multiple logistic regression within an expert-driven, spatially balanced, and information-theoretic framework. Our occurrence model incorporated forest structure variables that were derived from USFS Forest Inventory and Analysis plots and high-resolution satellite imagery. Results indicated that high canopy-bulk density, intermediate canopy-base heights, and low variation in tree density were strong predictors of territory occurrence. We used model-averaged parameter estimates for these variables to map and explore patterns of territory distribution across multiple land jurisdictions and ecological subregions. Our iterative modeling approach complements previous demographic studies in the region. It also provides a robust framework for integrating species conservation and landscape management in ongoing and future regional planning efforts.     

An integration of habitat evaluation,individual based modeling,and graph theory for a potential black bear population recovery in southeastern Texas,USA

Population recovery is difficult for species that require large contiguous areas of habitat, particularly within areas of heterogeneous land ownerships. Ecologically, potential for recovery success requires assessment of quantity, quality, and distribution of available habitat. Our objective was to evaluate habitat for a possible Louisiana black bear recovery in southeastern Texas. First, we categorized land cover and identified remote areas of highly suitable habitat. Next, we used the individual based simulation model J-walk to estimate ability of female black bears to move among remote habitat patches. Then, we applied graph theory to J-walk output to evaluate overall connectivity of remote habitat. An estimated 225,626 ha of remote habitat were identified in 901 patches, most of which was located within the eastern half of the study area. Network analysis showed specific areas where targeted conservation efforts may help black bear population expansion throughout the study region. Ultimately, enough habitat area exists to sustain a black bear population and it is best connected among public and private lands largely within the eastern half of the study area. Habitat evaluation will need to be revisited if black bears establish themselves locally and actual habitat use data become available. Regardless, our analysis demonstrates an important first step that may be incorporated into a larger adaptive management framework, updated, and replicated as more-detailed habitat suitability and land use data are available.