Spatial graphs as templates for habitat networks in boreal landscapes

Network topology serves as a useful model for biological systems at various scales. Contrary to many biological systems, spatial reference is crucial for habitat networks. Boreal forest landscapes provide a wide gradient of spatial patterns and, thus, unique network structures. Assuming forest-dwelling organisms in general aim to minimize travel distances during foraging, dispersal, etc., linear links across the landscape matrix constitute expected movement routes among forested areas in boreal landscapes. We quantified the number and length of links in a set of 57 boreal forest landscapes for four hierarchically nested graphs in order to compare the incremental changes in characteristics of resulting graph measures. The forest cover graphs consisted of the same set of forest patches, and hierarchical link types extracted from real landscapes: nearest neighbour graph (NN), minimum spanning tree (MST), Gabriel graph (GG) and minimum planar graph (MPG). Most of the links in graphs were NN and GG links. Commonly links were 100–200?m in length, but link lengths particularly in the GG and MPG shorten when the proportion of forest in landscapes increased. Most nodes had 3–5 links each, but the number of links per node depended on node size and the proportion of forest cover. GG and MPG graphs retain the topology of the underlying node layout. Changes in node pattern alter the NN and MST graphs more than GG and MPG. Variation in regional network topologies is likely to affect connectivity patterns in a landscape and, thus, many ecological processes that occur at a local scale. An appropriate network analysis enables the discovery and comparison of distinctive network patterns. Understanding network topologies provide practical tools for land use planning and biodiversity management of broader areas that target functional habitat networks.     

Fine-scale movement behaviors of calopterygid damselflies are influenced by landscape structure: an experimental manipulation

We explore the effect of differences in landscape structure, arising from habitat loss, on the fine-scale movement behaviors of two congeneric damselflies – Calopteryx aequabilis and C. maculata . Both species require streams for breeding and naiad development and both often use forest for foraging. We compare movement behaviors across three types of landscape: forested landscapes, where stream and forest habitat are adjacent; partially forested landscapes, where streams and forest habitat are disjunct, and non-forested landscapes, where little to no forest habitat is available. We employ a reciprocal transplant experiment to determine the extent to which movement along and away from streams is influenced by landscape structure and historical behavior or morphological adaptations. For both species, we show that both the propensity to move away from streams and rates of net displacement differ among landscape types. Both species move away from streams on landscapes with high or moderate levels of forest cover but neither moves away from streams on landscapes with little or no forest. Furthermore, C. maculata native to predominantly forested landscapes are more likely to move away from streams, regardless of the landscape structure they encounter, than are individuals native to moderately forested or non-forested landscapes. There was no effect of natal landscape on C. aequabilis . Comparisons with microlandscape studies suggest that there may be some general similarities among the different systems but these are clouded by uncertainty regarding the similarity of the underlying processes responsible for observed behavioral responses to landscape structure. Despite this uncertainty, animal movement behaviors are contingent upon the structure of the broader landscape, regardless of the absolute scale of the landscape.     

The influence of forest fragmentation and landscape pattern on American martens

1. We investigated the effects of forest fragmentation on American martens ( Martes americana Rhoads) by evaluating differences in marten capture rates (excluding recaptures) in 18 study sites with different levels of fragmentation resulting from timber harvest clearcuts and natural openings. We focused on low levels of fragmentation, where forest connectivity was maintained and non-forest cover ranged from 2% to 42%.
2. Martens appeared to respond negatively to low levels of habitat fragmentation, based on the significant decrease in capture rates within the series of increasingly fragmented landscapes. Martens were nearly absent from landscapes having > 25% non-forest cover, even though forest connectivity was still present.
3. Marten capture rates were negatively correlated with increasing proximity of open areas and increasing extent of high-contrast edges. Forested landscapes appeared unsuitable for martens when the average nearest-neighbour distance between open (non-forested) patches was <100 m. In these landscapes, the proximity of open areas created strips of forest edge and eliminated nearly all forest interior.
4. Small mammal densities were significantly higher in clearcuts than in forests, but marten captures were not correlated with prey abundance or biomass associated with clearcuts.
5. Conservation efforts for the marten must consider not only the structural aspects of mature forests, but the landscape pattern in which the forest occurs. We recommend that the combination of timber harvests and natural openings comprise <25% of landscapes ≥9 km² in size.
6. The spatial pattern of open areas is important as well, because small, dispersed openings result in less forest interior habitat than one large opening at the same percentage of fragmentation. Progressive cutting from a single patch would retain the largest amount of interior forest habitat.     

Long-term persistence of midsized to large-bodied mammals in Amazonian landscapes under varying contexts of forest cover

Both forest fragmentation and overhunting have profound effects on the structure of large-vertebrate assemblages in neotropical forests. However, the long-term value of habitat fragments for forest mammals remains poorly understood and few regional scale studies have replicated sampling across spatially independent landscapes. Here, we assess the species occupancy and abundance of midsized to large-bodied mammals within three neighbouring Amazonian forest landscapes varying widely in extent of forest cover. One of these consisted of forest fragments surrounded by semi-natural scrub savannahs that had been occupied by paleoindian populations for at least 7,000 years, whereas forest cover in the other two landscapes was either variegated or continuous. Data on species occurrence and abundance from diurnal and nocturnal line-transect surveys and local interviews in each landscape were used to examine the effects of forest cover and hunting pressure on mammal persistence within forest patches. The extent of forest cover was a key determinant of species persistence across the three landscapes, but populations of large-bodied species were either reduced or driven to local extinction by hunting even in the most forested and least fragmented landscape. Many game and non-game species persisted in forest isolates, even though, individually, these were likely too small to support viable populations. This study indicates that even small, long-term forest fragments may retain significant conservation value if they can be managed within the context of enhanced connectivity across wider fragmented landscapes.     

Habitat selection by capercaillie in summer and autumn: Is bilberry important?

The use of habitat by female and male adult capercaillie Tetrao urogallus during summer and autumn was studied by comparing the distribution of radio locations of birds with the availability of habitat at forest stand, home range and landscape level in an area of the Bavarian Alps, Germany. Capercaillie preferred forests with structural features typical of their main distribution range, the boreal forest: they selected large patches of old forest with moderate canopy cover of about 50%, and a well developed field layer with high proportions of bilberry Vaccinium myrtillus. Hens selected both home ranges and sites within home ranges in old forest. Ranges selected by cocks did not differ from availability in the study area, but they preferred old forest within their ranges. The size of home ranges was negatively related to bilberry cover both in hens and cocks. The distribution of bilberry also determined habitat use by capercaillie at the landscape scale. The study demonstrated that bilberry is the major determinant of the selection of habitat by capercaillie in landscapes with sparse and fragmentary cover of ericaceous shrubs, such as central Europe.     

Designing optimal human‐modified landscapes for forest biodiversity conservation

Agriculture and development transform forest ecosystems to human‐modified landscapes. Decades of research in ecology have generated myriad concepts for the appropriate management of these landscapes. Yet, these concepts are often contradictory and apply at different spatial scales, making the design of biodiversity‐friendly landscapes challenging. Here, we combine concepts with empirical support to design optimal landscape scenarios for forest‐dwelling species. The supported concepts indicate that appropriately sized landscapes should contain ≥ 40% forest cover, although higher percentages are likely needed in the tropics. Forest cover should be configured with c. 10% in a very large forest patch, and the remaining 30% in many evenly dispersed smaller patches and semi‐natural treed elements (e.g. vegetation corridors). Importantly, the patches should be embedded in a high‐quality matrix. The proposed landscape scenarios represent an optimal compromise between delivery of goods and services to humans and preserving most forest wildlife, and can therefore guide forest preservation and restoration strategies.     

Landscape ecology of boreal forests

It is becoming increasingly clear that densities and dynamics of various organisms often cannot be understood from the processes occurring within separate habitat patches. Effects from surrounding areas also have to be considered; a landscape perspective has to be employed. Habitat mosaics affect diversity and dynamics both in pristine and managed boreal landscapes. There is increasing interest in the landscape ecology of boreal forests and many important processes have been identified as being driven or affected by environmental heterogeneity. Comparisons between ecological processes in original and managed boreal landscapes are urgently needed.     

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.     

Metapopulation dynamics of a beetle species confined to burned forest sites in a managed forest region

Despite increasing awareness of the theoretical importance of habitat dynamics on metapopulations, only a few empirical studies have been conducted. We aimed to increase our understanding of how patch size, dynamics and connectivity affect colonization–extinction dynamics and the occurrence patterns of a beetle (Stephanopachys linearis), which breeds only in burned trees, existing as dynamic habitat patches that have become rare in managed forest landscapes. We assessed species’ presence/absence twice in all known habitat patches (i.e. > 1 ha sites where forest fires had occurred during the previous 2–15 yr) in a 200 × 150 km region of central Sweden, dominated by managed boreal forest. Evaluated over six years, the colonization rate was 47% and the local extinction risk was 65%. Probability of colonization increased with patch size (number of suitable trees in a site) and connectivity to occupied patches within 30 km, and decreased with increasing time since fire. Local extinction risk decreased with habitat patch size but increased, unexpectedly, with connectivity. Occurrence increased with patch size and decreased with increasing time since fire. At a regional scale, S. linearis tracks the fire dynamics by colonising sites with burned trees and by becoming extinct at rates which make the species rare at sites where burnt trees are more than eight years old. In managed boreal forest landscapes, a large proportion of sites may be created by prescribed burning (in our study area: 82%), and consequently human decisions strongly affect the future amount of habitat for fire‐dependent species and its spatial distribution. Stephanopachys linearis uses burned sites more often if more trees are retained and, to some extent, if sites are concentrated in those parts of a region that already support high population densities of the species.     

Forest landscape pattern in the KwaZulu–Natal midlands,South Africa: 50 years of change or stasis?

Abstract Understanding patterns and processes of habitat change is essential for managing and conserving forest fragments in anthropogenically altered landscapes. Digitized aerial photographs from 1944 and 1996 were examined for changes to the indigenous forest landscape in the Karkloof‐Balgowan archipelago in KwaZulu–Natal, South Africa. Attributes relating to proximate land‐use, patch shape, isolation and position in the landscape were used to determine putative causes of forest change. The total change in forest area was ?5.7% (forest covered 6739 ha in 1996). This is contrasted with previous reports for the period 1880–1940 that estimated change in total forest area of up to ?80%. Attrition was the predominant process of forest transformation between 1944 and 1996. Despite little overall change in forest area, 786 mostly small (<0.5 ha) forest patches were lost from the landscape, leaving 1277 forest patches in 1996. An increase in patch isolation, but no change in patch cohesion accompanied the changes in forest area. Ignoring patches that were eliminated, 514 patches decreased in area. This was partly a function of patch size, but the conversion of natural grassland to commercial plantation forestry in the matrix also influenced forest decline. Their small size and irregular shape caused forest patches in the region to be vulnerable to edge effects. Core area declined in a negative exponential way with increasing edge width and the total area of edge habitat exceeded that of core habitat at an edge width of only 50 m. Nevertheless, total core area decreased by only 2% (65 ha) between 1944 and 1996 because most of the eliminated patches were small and contained no core area. The large Karkloof forest (1649 ha) is a conservation priority for forest interior species, but the ecological role and biodiversity value of small forest patches should not be overlooked.     

Continuum or discrete patch landscape models for savanna birds? Towards a pluralistic approach

Conceptualising landscapes as a mosaic of discrete habitat patches is fundamental to landscape ecology, metapopulation theory and conservation biology. An emerging question in ecology is: when is the discrete patch model more appropriate than alternative and conceptually appealing models such as the continuum model? There is limited empirical testing of the utility of alternative landscape models compared to the discrete patch model for a range of species. In this paper, we constructed three alternative sets of models for testing the effect of landscape structure on diversity and abundance of a suite of woodland birds in a savanna landscape of northern Australia: the null model (only site‐scale habitat variables, landscape context not important), the continuum model, and the discrete patch model. We utilised high‐spatial resolution satellite images to quantify spatial gradients in tree cover density (the continuum model), and to then aggregate the fine‐scale heterogeneity in tree cover into discrete patches of trees, with grass cover forming the “matrix” (the discrete patch‐model). We then evaluated the relative importance of the alternative models using generalised linear models and an information theoretic approach. We found that the importance of the models varied among species, with no single model dominant. Species that move between open grassy areas and woody shelter responded well to the continuum model, reflecting the importance of gradients in density of forage (grasses) and cover (trees), while the discrete model performed best for species that forage in all vegetation strata, and nest predominantly in dense woody vegetation. This finding supports a pluralistic approach, highlighting the need for adopting and testing more than one landscape model in savanna landscapes, and in other landscapes that do not have a well defined patch structure.     

Isolation determines patterns of species presence in highly fragmented landscapes

In fragmented landscapes, changes in habitat availability, patch size, shape and isolation may affect survival of local populations. Proposing efficient conservation strategies for such species relies initially on distinguishing the particular effects of those factors. To address these issues, we investigated the occurrence of 3 bird species in fragmented Brazilian Atlantic Forest landscapes. Playback techniques were used to collect presence/absence data of these species inside 80 forest patches, and incidence models were used to infer their occupancy pattern from landscape spatial structure. The relative importance of patch size, shape and surrounding forest cover and isolation was assessed using a model selection approach based on maximum likelihood estimation. The presence of all species was in general positively affected by the amount of surrounding habitat and negatively affected by inter‐patch distances. The joint effects of patch size and the surrounding landscape characteristics were important determinants of occupancy for two species. The third species was affected only by forest cover and mean patch isolation. Our results suggest that local species presence is in general more influenced by the isolation from surrounding forests than by patch size alone. We found evidence that, in highly fragmented landscapes, birds that can not find patches large enough to settle may be able to overcome short distances through the matrix and include several nearby patches within their home‐ranges to complement their resource needs. In these cases, patches must be defined as functionally connected habitat networks rather than mere continuous forest segments. Bird conservation strategies in the Atlantic forest should focus on increasing patch density and connectivity, in order to implement forest networks that reduce the functional isolation between large remnants with remaining core habitat.     

Landscape attributes as drivers of the geographical variation in density of Sapajus nigritus Kerr, 1792, a primate endemic to the Atlantic Forest

Neotropical primates are among the most well studied forest mammals concerning their population densities. However, few studies have evaluated the factors that influence the spatial variation in the population density of primates, which limits the possibility of inferences towards this animal group, especially at the landscape-level. Here, we compiled density data of Sapajus nigritus from 21 forest patches of the Brazilian Atlantic Forest. We tested the effects of climatic variables (temperature, precipitation), landscape attributes (number of patches, mean inter-patch isolation distance, matrix modification index) and patch size on the population density using linear models and the Akaike information criterion. Our findings showed that the density of S. nigritus is influenced by landscape attributes, particularly by fragmentation and matrix modification. Overall, moderately fragmented landscapes and those surrounded by matrices with intermediate indexes of temporal modification (i.e., crop plantations, forestry) are related to high densities of this species. These results support the assumptions that ecologically flexible species respond positively to forest fragmentation. However, the non-linear relationship between S. nigritus density and number of patches suggests that even the species that are most tolerant to forest cover changes seem to respond positively only at an intermediate level of habitat fragmentation, being dependent of both a moderate degree of forest cover and a high quality matrix. The results we found here can be a common response to fragmentation for those forest dweller species that are able to use the matrix as complementary foraging sites.     

Avian fitness consequences match habitat selection at the nest‐site and landscape scale in agriculturally fragmented landscapes

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Importance of habitat patch size for occupancy and density of aspen-associated saproxylic beetles

In Fennoscandian boreal forests, aspen (Populus tremula) is one of the most important tree species for biodiversity. In this study we explore how occupancy and density of beetles associated with dead aspen are related to habitat patch size and connectedness in a 45,000 ha boreal managed forest landscape in central Sweden. Patch size was estimated as amount of breeding substrate and connectedness as crown cover of living aspen in the surrounding landscape. The beetles were sampled by sieving of bark or by inspection of species-characteristic galleries in 56 patches with dead aspen. Six of nine aspen-associated species (Xylotrechus rusticus, Ptilinus fuscus, Mycetophagus fulvicollis, Cyphaea curtula, Homalota plana and Endomychus coccineus) showed a positive significant relationship between habitat patch size and occupancy. For all these species, except C. curtula, there was also a significant positive relationship between patch size and density. Connectedness was not retained as a significant variable in the analyses. Species not defined as aspen-associated constituted a significantly larger proportion of the total density of individuals of saproxylic beetles in smaller habitat patches than in larger patches. Richness of aspen-associated species was positively related to habitat patch size. Efforts in the managed forest should be directed towards preserving and creating larger patches of living and dead aspen trees and increasing the amount of aspen at the landscape level.     

Forest bird diversity in Mediterranean areas affected by wildfires: a multi-scale approach

Fire is a key mechanism creating and maintaining habitat heterogeneity in Mediterranean landscapes by turning continuous woody landscapes into mosaics of forests and shrublands. Due to the long historical role of fires in the Mediterranean, we hypothesised a moderate negative effect of this type of perturbation on forest bird distribution at a landscape level. We conducted point bird censuses in Aleppo pine forest patches surrounded by burnt shrublands and studied the relationships between three ecological groups of bird species (forest canopy species, forest understorey species, and ubiquitous species) and the features of local habitat, whole patch and surrounding landscape. We used a multi-scale approach to assess the effects of landscape variables at increasing spatial scales on point bird richness. Regarding local habitat components, canopy species were positively associated with tall pines while understorey species with the cover of shrubs and plants from holm-oak forests. Forest birds were positively related to patch size and irregular forest shapes, that is, with high perimeter/size ratios. Thus, these species did not seem to perceive edges as low quality but rather favourable microhabitats. We did not detect any negative effect of isolation or cover of woodlands in the landscape on the presence of forest species after local habitat factors had been accounted for. Finally, only local habitat factors entered the model for ubiquitous species. We suggest that mosaic-like landscapes shaped by fires in the Mediterranean basin are not strongly associated with negative effects fragmentation on forest birds other than those related with habitat loss.     

Landscape connectivity dynamics based on network analysis in the Xishuangbanna Nature Reserve,China

Lack of landscape connectivity and habitat loss is major threats to biodiversity and ecosystem integrity in nature reserves aimed at conservation. In this study, we used structural pattern and functional connectivity metrics to analyze the spatial patterns and landscape connectivity of habitat patches for the Shangyong sub-reserve of the Xishuangbanna Nature Reserve from 1970, 1990, and 2000. On the basis of vegetation and land cover data, we applied the equivalent connected area ECA(PC) indicator to analyze the changes in forest connectivity. Four distance thresholds (2, 4, 8, 12 km) were considered to compare the patch importance of connectivity by dECA values. The results showed the declining trends of landscape connectivity measured by ECA(PC) index from 1970 to 2000. The importance of connectivity in each forest patch varied with the increment of dispersal distances at the patch level, and some important habitat patches, which exhibit a potential to enhance landscape connectivity, should be given more attention. The least-cost pathways based on network structure were displayed under four dispersal distances in three periods. The results showed that the number of paths among the fragments of forest patches exhibited radical increases for larger dispersal distances. Further correlation analyses of AWF, ECA (IIC), and ECA (PC) showed the weakest and least-frequent correlations with the structural pattern indices, while H presented more significant correlations with the PD fragmentation metric. Furthermore, Kendall's rank correlations between the forest patch area and functional connectivity indicators showed that dECA (PC) and dAWF indicators should provided the area-based prioritization of habitat patches. Moreover, the low-rank correlations showed that dF and dLCP can be considered as effective and appropriate indicators for the evaluation of habitat features and network patterns.     

Occupancy Models of Nesting-Season Habitat Associations of Red-Shouldered Hawks in Central Minnesota

ABSTRACT Red-shouldered hawks (Buteo lineatus) are a species of special conservation concern in much of the Great Lakes region, and apparent population declines are thought to be primarily due to habitat loss and alteration. To evaluate red-shouldered hawk-habitat associations during the nesting season and at the landscape scale, we conducted repeated call-broadcast surveys in central Minnesota, USA, across 3 landscapes that represented a range of landscape conditions as a result of differing management practices. In 2004, we conducted repeated call-broadcast surveys at 131 locations in 2 study areas, and in 2005, we surveyed 238 locations in 3 study areas. We developed models relating habitat characteristics at 2 spatial scales to red-shouldered hawk occupancy and assessed support for these models in an information-theoretic framework. Overall, a small proportion of nonforest (grass, clear-cut area, forest <5 yr old), and a large proportion of mature deciduous forest (>40 yr old), had the strongest association with red-shouldered hawk occupancy (proportion of sites occupied) at both spatial scales. The landscape conditions we examined appeared to contain a habitat transition important to red-shouldered hawks. We found, in predominately forest landscapes, the amount of open habitat was most strongly associated with red-shouldered hawk occupancy, but in landscapes that included slightly less mature forest and more extensive open habitats, the extent of mature deciduous forest was most strongly associated with red-shouldered hawk occupancy. Our results suggested that relatively small (<5 ha) patches of open habitat (clear-cuts) in otherwise forested landscapes did not appear to influence red-shouldered hawk occupancy. Whereas, in an otherwise similar landscape, with smaller amounts of mature deciduous forest and larger (>15 ha) patches of open habitat, red-shouldered hawk occupancy decreased, suggesting a threshold in landscape composition, based on both the amount of mature forest and open area, is important in managing forest landscapes for red-shouldered hawks. Our results show that during the nesting season, red-shouldered hawks in central Minnesota occupy at similar rates landscapes with different habitat compositions resulting from different management strategies and that management strategies that create small openings may not negatively affect red-shouldered hawk occupancy.     

Relationships between spatial configuration of tropical forest patches and woody plant diversity in northeastern Puerto Rico

The destruction and fragmentation of tropical forests are major sources of global biodiversity loss. A better understanding of anthropogenically altered landscapes and their relationships with species diversity and composition is needed in order to protect biodiversity in these environments. The spatial patterns of a landscape may control the ecological processes that shape species diversity and composition. However, there is little information about how plant diversity varies with the spatial configuration of forest patches especially in fragmented tropical habitats. The northeastern part of Puerto Rico provides the opportunity to study the relationships between species richness and composition of woody plants (shrubs and trees) and spatial variables [i.e., patch area and shape, patch isolation, connectivity, and distance to the Luquillo Experimental Forest (LEF)] in tropical forest patches that have regenerated from pasturelands. The spatial data were obtained from aerial color photographs from year 2000. Each photo interpretation was digitized into a GIS package, and 12 forest patches (24–34 years old) were selected within a study area of 28 km². The woody plant species composition of the patches was determined by a systematic floristic survey. The species diversity (Shannon index) and species richness of woody plants correlated positively with the area and the shape of the forest patch. Larger patches, and patches with more habitat edge or convolution, provided conditions for a higher diversity of woody plants. Moreover, the distance of the forest patches to the LEF, which is a source of propagules, correlated negatively with species richness. Plant species composition was also related to patch size and shape and distance to the LEF. These results indicate that there is a link between landscape structure and species diversity and composition and that patches that have similar area, shape, and distance to the LEF provide similar conditions for the existence of a particular plant community. In addition, forest patches that were closer together had more similarity in woody plant species composition than patches that were farther apart, suggesting that seed dispersal for some species is limited at the scale of 10 km.