Local habitat patch pattern of the Siberian flying squirrel in a managed boreal forest landscape

We examined how the structure of a boreal forest landscape is related to the occurrence of the Siberian flying squirrel Pteromys volans in northern Finland. The flying squirrel inhabits mature spruce-dominated ( Picea abies ) mixed forests and is categorised as vulnerable species due to habitat loss and change. We classified a landscape of 374.5 km² into potential habitat patches, potential dispersal areas, and areas incapable of being inhabited using national forest inventory data, and surveyed all 136 potential habitat patches for the presence of the species. Different landscape variables were defined, and also connections by the shortest distances to neighbouring habitat patches along both straight lines and least-cost distances based on specific movement costs were measured. Occupied patches were larger in size, contained more deciduous trees for food and nesting cavities, and were in closer proximity to the nearest occupied patches. Occupied patches were mainly located below 300 m a.s.l. The occurrence of flying squirrels was correctly predicted for 88% of the habitat patches using landscape variables. This modelling result proved to be rather general. In addition, the configuration of occupied patches was mainly clustered across the landscape, and distant occupied patches seemed to be linked to other patches via forested connections. We suggest that maintaining a clustered arrangement of good quality habitat patches and regenerating new potential habitat as well as dispersal areas between the habitat patches seem to be appropriate goals for long-term forest management planning to sustain populations of the flying squirrel in the landscape.     

Movements of the flying squirrel Pteromys volans in corridors and in matrix habitat

Elements of the landscape, such as patches of preferred habitat, matrix between patches, and corridors linking patches, differ as movement habitat for animals. To understand how landscape structure influences the movement and thus, population dynamics of animals, clear empirical knowledge on patterns of movement is needed. The Siberian flying squirrel inhabits spruce-dominated boreal forests from Finland to eastern Siberia. Numbers of flying squirrels have declined severely in Finland in past decades, probably due to modern forestry. We studied the movement of radio-collared adult flying squirrels in preferred (spruce forest) and in matrix habitat (open areas and other habitats with trees) in Finland 1997–2000, and determined whether the woodland strips connecting patches of preferred habitat could function as ecological corridors for flying squirrels.
Flying squirrels used woodland strips for inter-patch movements, but also used matrix with trees and were able to cross narrow open gaps. Males moved longer total distances and crossed edges more often than females. Males used matrix habitats for movement between spruce patches, and moved faster and more directly in the matrix than in the spruce forest. Females seldom changed spruce patches, but instead used the matrix for foraging. For both sexes probability of leaving the spruce forest patch correlated negatively with the size of the patch, but the type of connection the patch had to other patches did not affect the leaving probability. Due to efficient movement abilities of the flying squirrel and forest-dominated landscape structure of southern Finland, we suggest that conservation acts for maintaining viable populations of flying squirrels should focus on the quality of managed forest and the area of suitable breeding habitat (i.e. on habitat loss), but not necessarily on ecological corridors.     

Space use of the Siberian flying squirrel Ptevomys volans in fragmented forest landscapes

We studied the space use of the Siberian flying squirrel in fragmented forest landscapes in southern Finland in 1996- 1999. Twenty-four adult males and 23 adult females were radio trucked in three different study areas. The use of different landscape elements by flying squirrels was compared to their availability using compositional analysis. Effects of fragmentation on flying squirrels were studied by comparing the use of different habitat types, number of used patches, and size of home range to the principal components formed from the landscape variables patch size, patch isolation, area of spruce-dominated forests, and area of other habitats with trees.
Flying squirrels preferred spruce- and deciduous-dominated forests, but other habitats with trees were also used for moving and foraging. These less preferred habitats connected spruce and deciduous forest patches in a matrix of open areas. Males used several separate patches of preferred habitat, but females usually stayed in one patch. The spacing behaviour of males was affected by the structure of the landscape in a way that home ranges were larger and males used more spruce patches in the more highly fragmented landscape. Home-range size of females was not affected by the landscape structure. Basically, the space use of males seemed lo be reflected by the location and number of female home ranges. In the study area where female density was high, the home ranges of males were small. Our results give empirical evidence of the effects of habitat loss and landscape connectivity for the Siberian flying squirrel and the implications of these for the conservation of the species.     

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.     

Patch occupancy, population density and dynamics in a fragmented red squirrel Sciurusvulgaris population

We studied population dynamics of red squirrels in a group of small forest fragments, that cover only 6.5% of the total study area (4664 ha) and where distances to the nearest source population were up to 2.2 km. We tested effects of patch size, quality and isolation and supplementary feeding on patch occupation during 1995–99. Larger patches and patches with supplementary feeding had a higher probability of being occupied. No patch <3.5 ha was ever occupied. No effects of isolation were found, suggesting that the forest habitat in the study area is not sufficiently fragmented to influence red squirrel distribution across patches. For medium sized patches (3.7–21 ha), that were occupied some years, there was an increase in patch occupation over the years, even though overall population size tended to decrease. These patches had a high turnover, especially of males. Patches in which the squirrel population went extinct were recolonized within a year. For patches that were at least some years occupied, squirrel density depended on patch quality only. No effects of patch size, isolation and winter temperature on population density were found. These data suggest that in our study area habitat fragmentation has no effect on local squirrel density and that the random sample hypothesis explains the distribution pattern across patches.     

Forces structuring tree squirrel communities in landscapes fragmented by agriculture: species differences in perceptions of forest connectivity and carrying capacity

The North American red squirrel ( Tamiasciurus hudsonicus ) has expanded its range into the central hardwoods of the United States in conjunction with increasing forest fragmentation and declining gray squirrel ( Sciurus carolinensis ) populations. We used translocation experiments and patch occupancy data to test for interspecific differences in mobility and sensitivity to habitat loss and modification by agriculture. We released squirrels in fencerows to test the hypothesis that gray squirrels display inferior mobility relative to red and fox ( S. niger ) squirrels. Elapsed time to movement from fencerows for 76 individuals increased with distance to forest patches and harvesting of crops. Gray and red squirrels took longer to move from fencerows than fox squirrels, and gray squirrels were less successful at moving from fencerows than red and fox squirrels. Ecologically scaled landscape indices revealed the degree to which interspecific differences in mobility and individual area requirements accounted for the occurrence of these species across landscapes. Gray squirrel distribution was constrained both by individual area requirements and dispersal ability. Occurrence of red and fox squirrels was related to patch size but was unaffected by landscape connectivity.     

Landscape characteristics associated with the occurrence of the flying squirrel Pteromys volans in old-growth forests of northern Finland

We investigated whether the occurrence of the flying squirrel, a declined virgin forest species, was accounted for by the structure of the landscape in northeastern Finland We sampled 20 forest areas (1–77 km² in size) in 1995 for the occurrence of the species using a sampling plot method Landscape structure around occupied and unoccupied areas were compared using seven different radii ranging from 100 m lo 10 km Classified satellite images were analyzed with Geographic Information System (GIS) Ten areas could be determined to be occupied by the flying squirrel Results showed that occupied areas did not deviate from unoccupied ones in total area In unoccupied areas there were more mature pine-spruce forests, and less matrix habitat at the home range scale (100 m and 200 m radii) than in occupied areas Further more, in unoccupied forest areas open habitats (clear-cuts, open fens) were more common than in occupied ones at distances between 200 m and 2 km around sampling plots These differences resulted from both larger mean patch size of and smaller mean nearest neighbor distance between open habitat patches m unoccupied than in occupied forest areas According to a logistic regression analysis, information on the amount of open habitat within 1 km radius alone correctly classified 75% of the areas into occupied and unoccupied ones The probability of the presence of flying squirrel decreases with the increasing amount of open habitat within 1 km The results suggest that increasing the amount of open habitats (e g clear-cutting) at the local scale has a negative impact on the flying squirrel It is obvious that in the present situation where the population has severely declined and its optimal habitat is still diminishing, every patch suitable for the species - whether occupied or not -may be important for the population persistence     

Winter site occupancy patterns of the northern flying squirrel in boreal mixed-wood forests

We assessed habitat use by the northern flying squirrel (Glaucomys sabrinus) to test the hypothesis that the species is not a late-seral coniferous forest specialist in boreal mixedwood forests of northwestern Québec. We monitored 149 pairs of nest boxes over 3 visits during February–April 2008. A total of 31 pairs of nest boxes were occupied during the study. The average (3.7, 2.8, and 1.8) and maximum (8, 6, and 4) numbers of individuals in nest boxes decreased from the first to last visit. Average air temperatures were below freezing and increased with the arrival of spring (?23.9, ?14.5, and ?3.7 °C). Using GIS software, we created a 430-m (median dispersal distance) buffer around each pair of nest boxes and extracted landscape composition variables from digital forest inventory maps. We created models that potentially explained site occupancy by the northern flying squirrel as well as detection probability. Model selection indicated that the area of coniferous forests negatively affected site occupancy by flying squirrels while temperature positively influenced detection probability. Our results support the hypothesis that the northern flying squirrel is not a habitat specialist, but can occupy a wider range of forest cover types than previously recognized in various landscape contexts.     

Quantitative predictions for patch occupancy of capercaillie in fragmented habitats

A major conclusion of studying metapopulation biology is that species conservation should favor regional rather than local population persistence. Regional persistence is tightly linked to size, spatial configuration and quality of habitat patches. Hence it is important for the management of endangered species that priority patches can be identified. We developed a predictive model of patch occupancy by capercaillie, a threatened grouse species, based on a single snapshot of data. We used logistic regression to predict patch occupancy as a function of patch size, isolation, connectivity, relative altitude, and biogeographical area. The probability of a patch being occupied increased with patch size and increasing altitude, and decreased with increasing distance to the next occupied patch. Patch size was the most important predictor although occupied patches varied considerably in size. Our model only uses data on the number, size and spatial configuration of habitat patches. It is a useful tool to designate priority areas for conservation, i.e. large core patches with high resilience in habitat quality, smaller island‐patches that still have high probability of being inhabited or becoming recolonised, and patches functioning as “stepping stones”. If capercaillie is to be preserved, habitat suitability needs to be maintained in a functional network of patches that account for size and inter‐patch distance thresholds as found in this study. We suggest that similar area‐isolation relationships are valid for almost any region within the distribution range of capercaillie. The thresholds for occupancy are however likely to depend on characteristics of the respective landscape. The outcome of our study emphasises the need for future investigations that explore the relationship between patch occupancy, matrix quality and its resistance to dispersing individuals.     

The relative contribution of local habitat and landscape context to metapopulation processes: a dynamic occupancy modeling approach

Changes in site occupancy across habitat patches have often been attributed to landscape features in fragmented systems, particularly when considering metapopulations. However, failure to include habitat quality of individual patches can mask the relative importance of local scale features in determining distributional changes. We employed dynamic occupancy modeling to compare the strength of local habitat variables and metrics of landscape patterns as drivers of metapopulation dynamics for a vulnerable, high‐elevation species in a naturally fragmented landscape. Repeat surveys of Bicknell's thrush Catharus bicknelli presence/non‐detection were conducted at 88 sites across Vermont, USA in 2006 and 2007. We used an organism‐based approach, such that at each site we measured important local‐scale habitat characteristics and quantified landscape‐scale features using a predictive habitat model for this species. We performed a principal component analysis on both the local and landscape features to reduce dimensionality. We estimated site occupancy, colonization, and extinction probabilities while accounting for imperfect detection. Univariate, additive, and interaction models of local habitat and landscape context were ranked using AICc scores. Both local and landscape scales were important in determining changes in occupancy patterns. An interaction between scales was detected for occupancy dynamics indicating that the relationship of the parameters to local‐scale habitat conditions can change depending on the landscape context and vice versa. An increase in both landscape‐ and local‐scale habitat quality increased occupancy and colonization probability while decreasing extinction risk. Colonization and extinction were both more strongly influenced by local habitat quality relative to landscape patterns. We also identified clear, qualitative thresholds for landscape‐scale features. Conservation of large habitat patches in high‐cover landscapes will help ensure persistence of Bicknell's thrushes, but only if local scale habitat quality is maintained. Our results highlight the importance of incorporating information beyond landscape characteristics when investigating patch occupancy patterns in metapopulations.     

Role of the Siberian flying squirrel as an umbrella species for biodiversity in northern boreal forests

One of the potentially useful indirect shortcut methods in biodiversity conservation is the umbrella species concept. An umbrella species can be seen relatively demanding for the size of the area and probably also for certain habitat types: conservation management for the umbrella species would thus encompass other species preferring similar habitats but with smaller area requirements. As such, it has a comprehensive spatial aspect for landscape planning. We tested the role of the Siberian flying squirrel (Pteromys volans) as an umbrella species for wood dependent species among red-listed and old-growth forest associated polypores, epiphytic lichens and beetles. Flying squirrels inhabit home ranges of several to tens of hectares, and prefer mature spruce-dominated (Picea abies) mixed forests, which often have high amounts of dead wood. We carried out species surveys and trappings during 1 year from 20 mature spruce-dominated forest stands (altogether 162 ha), of which 12 were occupied by the flying squirrel. The amount of dead wood was higher in occupied stands than in unoccupied stands. We also found a tendency for a higher number of species and number of records in occupied stands, a relationship mostly due to the polypore species. The presence of the flying squirrel may reflect the habitat availability for species depending on dead and living wood, and assist in site selection of conservation areas. We suggest that the flying squirrel has potential as an umbrella species to partly enhance maintenance of biodiversity in northern boreal forests in Finland.     

Effects of Rain on Palm Springs Ground Squirrel Occupancy in the Sonoran Desert

ABSTRACT Long-term monitoring programs must use informative yet cost-effective methods. Occupancy estimates that incorporate detection probabilities are used with increasing frequency to describe species status and make management recommendations. Estimating changes in the occupancy of points over time in response to management actions or environmental changes may be especially useful for management of the Palm Springs round-tailed ground squirrel (Spermophilus tereticaudus chlorus), a subspecies covered under the Coachella Valley Multiple Species Habitat Conservation Plan and Natural Community Conservation Plan. In 2002 and 2003, we estimated occupancy and detection probability of ground squirrels across lands modeled as ground squirrel habitat by the Scientific Advisory Committee for the Habitat Conservation Plan and tested a priori hypotheses about how occupancy varied among vegetation and substrate types. In the 2003 study, we asked whether these associations were affected by winter rains after the 2002 drought year. Occupancy in 2003 was estimated at 0.99 (SE = 0.01) in Western honey mesquite (Prosopis glandulosa) on dunes and hummocks, and occupancy of the remaining modeled habitat was best described by distance to mesquite, with the occupancy probability decreasing with increasing distance from mesquite on dunes or hummocks. The best-supported model in 2002 described the distribution of ground squirrels as a function of only vegetation and substrate type. However, the best-supported models in 2003 suggested that distance to mesquite was a component of the occupancy of non-mesquite vegetation. Mesquite seems to provide high-quality habitat that can support ground squirrels at high occupancy probabilities that may breed successfully every year. In contrast, other vegetation types provide low-quality habitat that can only support ground squirrels at low occupancy probabilities that may only breed occasionally. Mesquite could be an essential refugium during drought years, and the 4 best-supported models in 2003 suggest that restoration of mesquite beginning near currently occupied mesquite patches could be critical for maintaining ground squirrel populations on the preserves.     

Natal habitat-biased dispersal in the Siberian flying squirrel

Theoretically, dispersers should target the habitat where prospects for fitness will be highest. Aiming for a habitat similar to the natal area (natal habitat-biased dispersal) has been hypothesized as a probable rule of thumb for dispersers, but has received very little empirical support to date. We investigated similarities between natal and post-dispersal settlement sites with radio-collared Siberian flying squirrels (Pteromys volans L.). Juveniles born in small patches and raised in nests close to patch edge settled in small patches and used nests close to edges after dispersal. In addition, post-dispersal use of dreys (versus cavities) was similar to that observed in natal sites. However, the quality of settlement habitat was unrelated to the quality of the natal site, which suggests that natal experience on average-quality habitats may not lead to ecological traps for flying squirrels. This study provides evidence that habitat selection at the landscape scale is influenced by habitat of natal area.     

Multilevel landscape utilization of the Siberian flying squirrel: Scale effects on species habitat use

Animals use and select habitat at multiple hierarchical levels and at different spatial scales within each level. Still, there is little knowledge on the scale effects at different spatial levels of species occupancy patterns. The objective of this study was to examine nonlinear effects and optimal‐scale landscape characteristics that affect occupancy of the Siberian flying squirrel, Pteromys volans, in South‐ and Mid‐Finland. We used presence–absence data (n = 10,032 plots of 9 ha) and novel approach to separate the effects on site‐, landscape‐, and regional‐level occupancy patterns. Our main results were: landscape variables predicted the placement of population patches at least twice as well as they predicted the occupancy of particular sites; the clear optimal value of preferred habitat cover for species landscape‐level abundance is a surprisingly low value (10% within a 4 km buffer); landscape metrics exert different effects on species occupancy and abundance in high versus low population density regions of our study area. We conclude that knowledge of regional variation in landscape utilization will be essential for successful conservation of the species. The results also support the view that large‐scale landscape variables have high predictive power in explaining species abundance. Our study demonstrates the complex response of species occurrence at different levels of population configuration on landscape structure. The study also highlights the need for data in large spatial scale to increase the precision of biodiversity mapping and prediction of future trends.     

Population consequences of movement decisions in a patchy landscape

Complex, human‐dominated landscapes provide unique challenges to animals. In landscapes fragmented by human activity, species whose home ranges ordinarily consist of continuous habitat in pristine environments may be forced to forage among multiple smaller habitat patches embedded in an inhospitable environment. Furthermore, foragers often must decide whether to traverse a heterogeneous suite of landscape elements that differ in risk of predation or energetic costs. We modeled population consequences of foraging decisions for animals occupying patches embedded in a heterogeneous landscape. In our simulations, animals were allowed to use three different rules for moving between patches: a) optimal selection resulting from always choosing the least‐cost path; b) random selection of a movement path; and c) probabilistic selection in which path choice was proportional to an animal's probability of survival while traversing the path. The resulting distribution of the population throughout the landscape was dependent on the movement rule used. Least‐cost movement rules (a) produced landscapes that contained the highest average density of consumers per patch. However, optimal movement resulted in an all‐or‐none pattern of occupancy and a coupling of occupied patches into pairs that effectively reduced the population to a set of sub‐populations. Random and probabilistic rules, (b and c), in relatively safe landscapes produced similar average densities and 100% occupancy of patches. However, as the level of risk associated with travel between patches increased, random movement resulted in an all‐or‐none occupancy pattern while occupied patches in probabilistic populations went extinct independently of the other patches. Our results demonstrate strong effects of inter‐patch heterogeneity and movement decisions on population dynamics, and suggest that models investigating the persistence of species in complex landscapes should take into account the effects of the intervening landscape on behavioral decisions affecting animal movements between patches.     

Heterogeneity in patch quality buffers metapopulations from pathogen impacts

Many wildlife species persist on a network of ephemerally occupied habitat patches connected by dispersal. Provisioning of food and other resources for conservation management or recreation is frequently used to improve local habitat quality and attract wildlife. Resource improvement can also facilitate local pathogen transmission, but the landscape-level consequences of provisioning for pathogen spread and habitat occupancy are poorly understood. Here, we develop a simple metapopulation model to investigate how heterogeneity in patch quality resulting from resource improvement influences long-term metapopulation occupancy in the presence of a virulent pathogen. We derive expressions for equilibrium host–pathogen outcomes in terms of provisioning effects on individual patches (through decreased patch extinction rates) and at the landscape level (the fraction of high-quality, provisioned patches), and highlight two cases of practical concern. First, if occupancy in the unprovisioned metapopulation is sufficiently low, a local maximum in occupancy occurs for mixtures of high- and low-quality patches, such that further increasing the number of high-quality patches both lowers occupancy and allows pathogen invasion. Second, if the pathogen persists in the unprovisioned metapopulation, further provisioning can result in all patches becoming infected and in a global minimum in occupancy. This work highlights the need for more empirical research on landscape-level impacts of local resource provisioning on pathogen dynamics.     

The larval ecology of the butterfly Euphydryas desfontainii (Lepidoptera: Nymphalidae) in SW-Portugal: food plant quantity and quality as main predictors of habitat quality

Corresponding to theory, the persistence of metapopulations in fragmented landscapes depends on the area of suitable habitat patches and their degree of isolation, mediating the individual exchange between habitats. More recently, habitat quality has been highlighted as being equally important. We therefore assess the role of habitat area, isolation and quality for the occupancy of larval stages of the regionally threatened butterfly Euphydryas desfontainii occurring in grassland habitats comprising the host plant Dipsascus comosus. We put a special focus on habitat quality which was determined on two spatial scales: the landscape (among patches) and the within-patch level. On the landscape level, occupancy of caterpillars was determined by a presence-absence analysis at 28 host plant patches. On the within-patch level, oviposition site selection was studied by comparing 159 host plants with egg clutches to a random sample of 253 unoccupied host plants within six habitat patches. The occupancy of caterpillars and presence of egg clutches on host plants was then related to several predictors such as patch size and isolation on the landscape level and host plant characteristics and immediate surroundings on the within patch level. On the landscape level, only host plant abundance was related to the presence of caterpillars, while size and isolation did not differ between occupied and unoccupied patches. However, the weak discrimination of larval stages among patches changed on the within-patch level: here, several microclimatic predictors such as sunshine hours and topography, host plant morphology and phenology as well as further potential host plants in the immediate surroundings of the plant chosen for oviposition strongly determined the presence of egg clutches. We strongly suggest promoting the presence of the host plant in topographically and structurally rich habitat patches to offer potential for microclimatic compensation for a species considered threatened by climate change.     

Thresholds of riparian forest use by terrestrial mammals in a fragmented Amazonian deforestation frontier

Species persistence in fragmented landscapes is intimately related to the quality, structure, and context of remaining habitat remnants. Riparian vegetation is legally protected within private landholdings in Brazil, so we quantitatively assessed occupancy patterns of terrestrial mammals in these remnants, examining under which circumstances different species effectively use them. We selected 38 riparian forest patches and five comparable riparian sites within continuous forest, at which we installed four to five camera-traps per site (199 camera-trap stations). Terrestrial mammal assemblages were sampled for 60 days per station during the dry seasons of 2013 and 2014. We modelled species occupancy and detection probabilities within riparian forest remnants, and examined the effects of patch size, habitat quality, and landscape structure on occupancy probabilities. We then scaled-up modelled occupancies to all 1915 riparian patches throughout the study region to identify which remnants retain the greatest potential to work as habitat for terrestrial vertebrates. Of the ten species for which occupancy was modelled, six responded to forest quality (remnant degradation, cattle intrusion, palm aggregations, and understorey density) or structure (remnant width, isolation, length, and area of the patch from which it originates). Patch suitability was lower considering habitat quality than landscape structure, and virtually all riparian remnants were unsuitable to maintain a high occupancy probability for all species that responded to forest patch quality or structure. Beyond safeguarding legal compliance concerning riparian remnant amount, ensuring terrestrial vertebrate persistence in fragmented landscapes will require curbing the drivers of forest degradation within private landholdings.     

Monitoring Golden-Cheeked Warblers on Private Lands in Texas

ABSTRACT A majority of North American breeding habitat for neotropical migrants exists on private lands, requiring monitoring strategies focused on habitat in these private holdings. We outline study designs and protocols using repeated Presence-Absence surveys across a gradient of patch sizes to develop a range-wide monitoring program for the endangered golden-cheeked warbler (Dendroica chrysoparia) in Texas, USA. We surveyed 200–400 point-count locations across approximately 30 private properties annually from 2005 to 2008. We used data from our surveyed patches (n = 147) and the Ψ (occupancy), p (detection), and γ = 1 - ɛ parameterization to estimate patch dynamics and associated detection probabilities for golden-cheeked warblers. Patch size had a strong association with patch occupancy, and all patches >160 ha were predicted to be occupied. We found no evidence that large golden-cheeked warbler populations located on public lands in the vicinity of our study area influenced occupancy dynamics. We conducted simulations across a range of detection probabilities to evaluate potential sample sizes for both standard- and removal-based occupancy modeling. Simulations using parameter estimates from our analysis indicated that removal-based sampling is superior to standard sampling. Based on our results, surveying golden-cheeked warbler presence in oak-juniper (Quercus-Juniperus) patches under a removal modeling framework should be considered as one alternative for range-wide monitoring programs because patch-level monitoring would be necessary to estimate proportion of range occupied. Large contiguous patches are rare across the species’ range; hence, conservation and management of the mosaic of smaller patches within a landscape context would be required for maintaining species viability. Thus, we recommend the identification of areas where smaller, contiguous patches represent a significant portion of the available habitat within the local landscape and targeting these areas for habitat maintenance and improvement.     

The roles of landscape context, niche breadth, and range boundaries in predicting species responses to habitat alteration

Extant species in human‐dominated landscapes differ in their sensitivity to habitat loss and fragmentation, although extinctions induced by environmental alteration reduce variation and result in a surviving subset of species with some degree of ‘resistance’. Here, we test the degree to which variable responses to habitat alteration are (1) essentially an inherent property of a taxon subject to constraints imposed by its geographical range, as suggested by Swihart et al. (2003), (2) a function of the landscape in which a species occurs, or (3) a function of spatial trends occurring on large scales. We used data collected on 33 vertebrate species during 2001–04 across the upper Wabash River basin, Indiana, in 35 square ‘landscapes’, each 23 km² in size. Six species of forest rodent, six species of grassland rodents, seven species of bats, eight species of aquatic turtles, and six species of amphibians were sampled at 504, 212, 590, 228, and 625 patches, respectively. The fraction of patches of primary habitat (e.g. forests for tree squirrels, wetlands for aquatic turtles) occupied by a target species was used as a response variable. On a basin‐wide scale, 47% of variation in proportional occupancy among species could be explained by taxon‐specific variables; occupancy rates were related positively to niche breadth and negatively to the proximity of a geographical range boundary. After controlling for species effects, landscape‐level occupancy rates varied significantly for 16 of 33 species, with variation partitioned among landscape variables alone (mean = 11% of variation), spatial trend variables alone (26%), and both variable sets jointly (8%). Among landscape variables, percentage forest cover positively affected occupancy rates of three bat species and a tree squirrel. Variation in occupancy rates among landscapes was consistent with large‐scale spatial trends for 13 species. Our findings demonstrate the general importance of niche breadth as a predictor of species responses to habitat alteration and highlight the importance of viewing the effects of habitat loss and fragmentation at multiple spatial scales.