The Viability of Metapopulations: Individual Dispersal Behaviour Matters

Metapopulation models are frequently used for analysing species–landscape interactions and their effect on structure and dynamic of populations in fragmented landscapes. They especially support a better understanding of the viability of metapopulations. In such models, the processes determining metapopulation viability are often modelled in a simple way. Animals’ dispersal between habitat fragments is mostly taken into account by using a simple dispersal function that assumes the underlying process of dispersal to be random movement. Species-specific dispersal behaviour such as a systematic search for habitat patches is likely to influence the viability of a metapopulation. Using a model for metapopulation viability analysis, we investigate whether such specific dispersal behaviour affects the predictions of ranking orders among alternative landscape configurations rated regarding their ability to carry viable metapopulations. To incorporate dispersal behaviour in the model, we use a submodel for the colonisation rates which allows different movement patterns to be considered (uncorrelated random walk, correlated random walk with various degrees of correlation, and loops). For each movement pattern, the landscape order is determined by comparing the resulting mean metapopulation lifetime T_m of different landscape configurations. Results show that landscape orders can change considerably between different movement patterns. We analyse whether and under what circumstances dispersal behaviour influences the ranking orders of landscapes. We find that the ‘competition between patches for migrants’ – i.e. the fact that dispersers immigrating into one patch are not longer available as colonisers for other patches – is an important factor driving the change in landscape ranks. The implications of our results for metapopulation modelling, planning and conservation are discussed.     

What you see is where you go? Modeling dispersal in mountainous landscapes

Inter-patch connectivity can be strongly influenced by topography and matrix heterogeneity, particularly when dealing with species with high cognitive abilities. To estimate dispersal in such systems, simulation models need to incorporate a behavioral component of matrix effects to result in more realistic connectivity measures. Inter-patch dispersal is important for the persistence of capercaillie (Tetrao urogallus) in central Europe, where this endangered grouse species lives in patchy populations embedded in a mountainous landscape. We simulated capercaillie movements with an individual-based, spatially explicit dispersal model (IBM) and compared the resulting connectivity measure with distance and an expert estimation. We used a landscape comprising discrete habitat patches, temporary habitat, non-habitat forests, and non-habitat open land. First, we assumed that dispersing individuals have perfect knowledge of habitat cells within the perceptual range (null model). Then, we included constraints to perception and accessibility, i.e., mountain chains, open area and valleys (three sub-models). In a full model, all sub-models were included at once. Correlations between the different connectivity measures were high (Spearman’s ρ > 0.7) and connectivity based on the full IBM was closer to expert estimation than distance. For selected cases, simple distance differed strongly from the full IBM measure and the expert estimation. Connectivity based on the IBM was strongly sensitive to the size of perceptual range with higher sensitivity for the null model compared to the full model that included context dependent perceptual ranges. Our heuristic approach is adequate for simulating movements of species with high cognitive abilities in strongly structured landscapes that influence perception and permeability.     

Animal movements and population dynamics in heterogeneous landscapes

Organisms respond to environmental heterogeneity at different scales and in different ways. These differences are consequences of how the movement characteristics of animals—their movement rates, directionality, turning frequencies, and turning angles—interact with patch and boundary features in landscape mosaics. The interactions of movement patterns with landscape features in turn produce spatial patterns in individual space-use, population dynamics and dispersion, gene flow, and the redistribution of nutrients and other materials. We describe several theoretical approaches for modeling the diffusion, foraging behavior, and population dynamics of animals in heterogeneous landscapes, including: (1) scaling relationships derived from percolation theory and fractal geometry, (2) extensions of traditional patch-based metapopulation models, and (3) individual-based, spatially explicit models governed by local rules. We conclude by emphasizing the need to couple theoretical models with empirical studies and the usefulness of ‘microlandscape’ investigations.     

Genetic isolation by distance and landscape connectivity in the American marten (Martes americana)

Empirical studies of landscape connectivity are limited by the difficulty of directly measuring animal movement. ‘Indirect’ approaches involving genetic analyses provide a complementary tool to ‘direct’ methods such as capture–recapture or radio-tracking. Here the effect of landscape on dispersal was investigated in a forest-dwelling species, the American marten (Martes americana) using the genetic model of isolation by distance (IBD). This model assumes isotropic dispersal in a homogeneous environment and is characterized by increasing genetic differentiation among individuals separated by increasing geographic distances. The effect of landscape features on this genetic pattern was used to test for a departure from spatially homogeneous dispersal. This study was conducted on two populations in homogeneous vs. heterogeneous habitat in a harvested boreal forest in Ontario (Canada). A pattern of IBD was evidenced in the homogeneous landscape whereas no such pattern was found in the near-by harvested forest. To test whether landscape structure may be accountable for this difference, we used effective distances that take into account the effect of landscape features on marten movement instead of Euclidean distances in the model of isolation by distance. Effective distances computed using least-cost modeling were better correlated to genetic distances in both landscapes, thereby showing that the interaction between landscape features and dispersal in Martes americana may be detected through individual-based analyses of spatial genetic structure. However, the simplifying assumptions of genetic models and the low proportions in genetic differentiation explained by these models may limit their utility in quantifying the effect of landscape structure.     

Integrative approach for landscape-based graph connectivity analysis: a case study with the common frog (Rana temporaria) in human-dominated landscapes

Graph-based analysis is a promising approach for analyzing the functional and structural connectivity of landscapes. In human-shaped landscapes, species have become vulnerable to land degradation and connectivity loss between habitat patches. Movement across the landscape is a key process for species survival that needs to be further investigated for heterogeneous human-dominated landscapes. The common frog (Rana temporaria) was used as a case study to explore and provide a graph connectivity analysis framework that integrates habitat suitability and dispersal responses to landscape permeability. The main habitat patches influencing habitat availability and connectivity were highlighted by using the software Conefor Sensinode 2.2. One of the main advantages of the presented graph-theoretical approach is its ability to provide a large choice of variables to be used based on the study’s assumptions and knowledge about target species. Based on dispersal simulation modelling in potential suitable habitat corridors, three distinct patterns of nodes connections of differing importance were revealed. These patterns are locally influenced by anthropogenic barriers, landscape permeability, and habitat suitability. And they are affected by different suitability and availability gradients to maximize the best possible settlement by the common frog within a terrestrial habitat continuum. The study determined the key role of landscape-based approaches for identifying the “availability-suitability-connectivity” patterns from a local to regional approach to provide an operational tool for landscape planning.     

Connectivity of agroecosystems: dispersal costs can vary among crops

Knowledge of how habitat heterogeneity affects dispersal is critical for conserving connectivity in current and changing landscapes. However, we generally lack an understanding of how dispersal costs and animal movements vary among crops characteristic of agroecosystems. We hypothesized that a physiological constraint, desiccation risk, influences movement behavior among crops and other matrix habitats (corn, soybean, forest, prairie) in Ambystoma tigrinum (tiger salamander) in Illinois, USA. In a desiccation experiment, salamanders were added to enclosures in four replicate plots of each matrix habitat, and water loss was measured every 12 h for 48 h. Changes in water loss were examined using a linear mixed model. Water loss varied among treatments, over time, and there was a significant treatment-time interaction. Water loss was greater in corn and prairie than in forest and soybean. To assess whether salamanders move through matrix habitats that minimize desiccation, we tracked movements of individuals released on edges between habitats for two treatment combinations: soybean–corn, and soybean–prairie. As predicted based on our desiccation experiment, movements were oriented towards soybean in both cases. Thus, variation in desiccation risk among matrix habitats likely influenced movement decisions by salamanders, although other factors such as predation risk could have contributed to habitat choice. We argue that conceptualizing dispersal cost as uniformly high in all crop types is too simplistic. Estimating crop-specific dispersal costs and movement patterns may be necessary for constructing effective measures of landscape connectivity in agroecosystems.     

Movements of cactus bugs: Patch transfers,matrix resistance,and edge permeability

Individual movement is a key process affecting the distribution of animals in heterogeneous landscapes. For specialist species in patchy habitat, a central issue is how dispersal distances are related to landscape structure. We compared dispersal distances for cactus bugs (Chelinidea vittiger) on two naturally fragmented landscapes (≤ 4% suitable habitat) with different matrix structures (i.e., vegetation height of nonsuitable habitat between suitable patches). Using mark-release-recapture studies, we determined that most transfers between cactus patches occurred during the mating season. Dispersal distances were reduced by > 50% on the landscape that had reduced structural connectivity due to relatively high matrix structure and low patch density. An experiment with detailed movement pathways demonstrated that greater matrix structure decreased mean step lengths, reduced directionality, and thus decreased net displacement by > 60%. However, habitat edges between two matrix elements that differed substantially in resistance to movement were completely permeable. Therefore, the difference in distributions of dispersal distances between the two landscapes mainly reflected the average resistance of matrix habitat and not the level of matrix heterogeneity per se. Our study highlights the merits of combining estimates of dispersal distances with insights on mechanisms from detailed movement pathways, and emphasizes the difficulty of treating dispersal distances of species as fixed traits independent of landscape structure.     

Dispersal success on fractal landscapes: a consequence of lacunarity thresholds

Habitat fragmentation is expected to disrupt dispersal, and thus we explored how patch metrics of landscape structure, such as percolation thresholds used to define landscape connectivity, corresponded with dispersal success on neutral landscapes. We simulated dispersal as either a purely random process (random direction and random step lengths) or as an area-limited random walk (random direction, but movement limited to an adjacent cell at each dispersal step) and quantified dispersal success for 1000 individuals on random and fractal landscape maps across a range of habitat abundance and fragmentation. Dispersal success increased with the number of cells a disperser could search (m), but poor dispersers (m<5) searching via area-limited dispersal on fractal landscapes were more successful at locating suitable habitat than random dispersers on either random or fractal landscapes. Dispersal success was enhanced on fractal landscapes relative to random ones because of the greater spatial contagion of habitat. Dispersal success decreased proportionate to habitat loss for poor dispersers (m=1) on random landscapes, but exhibited an abrupt threshold at low levels of habitat abundance (p<0.1) for area-limited dispersers (m<10) on fractal landscapes. Conventional metrics of patch structure, including percolation, did not exhibit threshold behavior in the region of the dispersal threshold. A lacunarity analysis of the gap structure of landscape patterns, however, revealed a strong threshold in the variability of gap sizes at low levels of habitat abundance (p<0.1) in fractal landscapes, the same region in which abrupt declines in dispersal success were observed. The interpatch distances or gaps across which dispersers must move in search of suitable habitat should influence dispersal success, and our results suggest that there is a critical gap-size structure to fractal landscapes that interferes with the ability of dispersers to locate suitable habitat when habitat is rare. We suggest that the gap structure of landscapes is a more important determinant of dispersal than patch structure, although both are ultimately required to predict the ecological consequences of habitat fragmentation.     

Modeling patch occupancy: Relative performance of ecologically scaled landscape indices

In fragmented landscapes, the likelihood that a species occupies a particular habitat patch is thought to be a function of both patch area and patch isolation. Ecologically scaled landscape indices (ESLIs) combine a species’ ecological profile, i.e., area requirements and dispersal ability, with indices of patch area and connectivity. Since their introduction, ESLIs for area have been modified to incorporate patch quality. ESLIs for connectivity have been modified to incorporate niche breadth, which may influence a species’ ease in crossing the non-habitat matrix between patches. We evaluated the ability of 4 ESLIs, the original and modified indices of area and connectivity, to explain patterns in patch occupancy of 5 forest rodents. Occupancy of eastern gray squirrels (Sciurus carolinensis), North American red squirrels (Tamiasciurus hudsconicus), fox squirrels (Sciurus niger), white-footed mice (Peromyscus leucopus), and eastern chipmunks (Tamias striatus) was modeled at 471 sites in 35 landscapes sampled from the upper Wabash River basin in Indiana. Models containing ESLIs received support for gray squirrels, red squirrels, and chipmunks. Modified ESLIs were important in models for red squirrels. However, none of the models demonstrated high predictive ability. Incorporating habitat quality and using surrogate measures of dispersal can have important effects on model results. Additionally, different responses of species to area, isolation, and habitat quality suggest that generalizing patterns of metapopulation dynamics was not justified, even across closely related species.     

Characterizing Fine-scale Patterns of Alternative Agricultural Landscapes with Landscape Pattern Indices

Landscape pattern indices are common tools of landscape ecologists, affording comparisons of different study areas, or the same study area at different times. Since the advent of popular index-calculating software, more landscapes can be analyzed in short amounts of time, yet the behaviour of landscape pattern indices can vary for different contexts or data characteristics, complicating interpretation. I applied a selected set of landscape pattern indices to fine-resolution (3 m) data representing a highly fragmented landscape – Corn Belt Iowa agriculture – to investigate the performance of landscape pattern indices. Indices measured pattern attributes that affect the viability of small mammal populations, namely habitat proportion and connectivity and landscape grain size and heterogeneity. Results showed that the performance of indices for fine-resolution data can be highly variable, depending upon data and contextual issues like the presence of linear elements and the amount of habitat. For these Corn Belt landscapes good habitat proportions and patch sizes were small (commonly less than 10% and less than 1 ha, respectively), and connectivity was variable depending on the measure. Aggregation and mean nearest neighbour indices performed better than other connectivity indices. Fine-resolution data representing highly fragmented landscapes can raise difficulties for indices of landscape configuration. Landscape pattern indices require improvement to perform better for increasingly available fine-resolution data representing common landscape types.     

Landscape – What’s in it? Trends in European Landscape Science and Priority Themes for Concerted Research

Reflecting on the other papers in this special issue, this synopsis characterises some essential trends in European Landscape Ecology, including the challenges it is facing in society. It describes the various perspectives on the ‘contents’ of landscape that are currently being practiced, and especially considers the notion of ‘environment’ as something intrinsic to human activity. Landscape classification and typology are discussed in their potential but limited use for landscape science. The specificity of the European approach appears to be related to the large diversity of cultural landscapes, currently losing their functional ties with the land-use systems that had formed them. European landscape research reports show a large commitment to this decreasing diversity, a dedication characterised by a strong sense of ‘loss and grief’. On the other hand, it is concluded that European landscape research has a specific niche with a clear focus on applied landscape studies explicitly including people’s perceptions and images, as well as the participation of the public and stakeholders. Since globalisation tends to reinforce the detachment of people from their environment; an increased effort is needed to compensate for this effect, and therefore the consideration of the various dimensions of the landscape is today more pertinent than ever. Meeting the challenges of present landscapes, in the face of new multifunctional demands in old diverse landscapes, requires more than before the combination of various perspectives and methods, and of various scales of application, in order to design innovative and adaptive paths for the future.     

Landscape resistance and American marten gene flow

Landscape heterogeneity can influence animal dispersal by causing a directional bias in dispersal rate, as certain landscape configurations might promote, impede, or prevent movement and gene flow. In forested landscapes, logging operations often contribute to heterogeneity that can reduce functional connectivity for some species. American martens (Martes americana) are one such species, as they are considered specialists of late-seral coniferous forests. We assessed marten gene flow to test the hypothesis that habitat management has maintained landscape connectivity for martens in the managed forests of Ontario, Canada. We genotyped 653 martens at 12 microsatellite loci, sampled from 29 sites across Ontario. We expected that if forest management has an effect on marten gene flow, we would see a correlation between effective resistance, estimated by circuit theory, and genetic distance, estimated by population graphs. Although we found a positive relationship between effective resistance and genetic distance (Mantel r = 0.249, P < 0.001), marten gene flow was better described by isolation by Euclidean distance (Mantel r = 0.410, P < 0.001). Our results suggest that managed forests in Ontario are well connected for marten and neither impede nor promote marten gene flow at the provincial scale.     

Making molehills out of mountains: landscape genetics of the Mojave desert tortoise

Heterogeneity in habitat often influences how organisms traverse the landscape matrix that connects populations. Understanding landscape connectivity is important to determine the ecological processes that influence those movements, which lead to evolutionary change due to gene flow. Here, we used landscape genetics and statistical models to evaluate hypotheses that could explain isolation among locations of the threatened Mojave desert tortoise (Gopherus agassizii). Within a causal modeling framework, we investigated three factors that can influence landscape connectivity: geographic distance, barriers to dispersal, and landscape friction. A statistical model of habitat suitability for the Mojave desert tortoise, based on topography, vegetation, and climate variables, was used as a proxy for landscape friction and barriers to dispersal. We quantified landscape friction with least-cost distances and with resistance distances among sampling locations. A set of diagnostic partial Mantel tests statistically separated the hypotheses of potential causes of genetic isolation. The best-supported model varied depending upon how landscape friction was quantified. Patterns of genetic structure were related to a combination of geographic distance and barriers as defined by least-cost distances, suggesting that mountain ranges and extremely low-elevation valleys influence connectivity at the regional scale beyond the tortoises’ ability to disperse. However, geographic distance was the only influence detected using resistance distances, which we attributed to fundamental differences between the two ways of quantifying friction. Landscape friction, as we measured it, did not influence the observed patterns of genetic distances using either quantification. Barriers and distance may be more valuable predictors of observed population structure for species like the desert tortoise, which has high dispersal capability and a long generation time.     

Independent effects of connectivity predict homing success by northern flying squirrel in a forest mosaic

Landscape composition and configuration, often termed as habitat loss and fragmentation, are predicted to reduce species population viability, partly due to the restriction of movement in the landscape. Unfortunately, measuring the effects of habitat loss and fragmentation on functional connectivity is challenging because these variables are confounded, and often the motivation for movement by target species is unknown. Our objective was to determine the independent effects of landscape connectivity from the perspective of a mature forest specialist—the northern flying squirrel (Glaucomys sabrinus). To standardize movement motivation, we translocated 119 squirrels, at varying distances (0.18–3.8 km) from their home range across landscapes representing gradients in both habitat loss and fragmentation. We measured the physical connectedness of mature forest using an index of connectivity (landscape coincidence probability). Patches were considered connected if they were within the mean gliding distance of a flying squirrel. Homing success increased in landscapes with a higher connectivity index. However, homing time was not strongly predicted by habitat amount, connectivity index, or mean nearest neighbour and was best explained as a simple function of sex and distance translocated. Our study shows support for the independent effects of landscape configuration on animal movement at a spatial scale that encompasses several home ranges. We conclude that connectivity of mature forest should be considered for the conservation of some mature forest specialists, even in forest mosaics where the distinction between habitat and movement corridors are less distinct.     

Textural Ordination Based on Fourier Spectral Decomposition: A Method to Analyze and Compare Landscape Patterns

We propose an approach to texture characterization and comparison that directly uses the information of digital images of the earth surface without requesting a prior distinction of structural ‘patches’. Digital images are partitioned into square ‘windows’ that define the scale of the analysis and which are submitted to the two-dimensional Fourier transform for extraction of a simplified textural characterization (in terms of coarseness) via the computation of a ‘radial’ power spectrum. Spectra computed from many images of the same size are systematically compared by means of a principal component analysis (PCA), which provides an ordination along a limited number of coarseness vs. fineness gradients. As an illustration, we applied this approach to digitized panchromatic air photos depicting various types of land cover in a semiarid landscape of northern Cameroon. We performed ‘textural ordinations’ at several scales by using square windows with sides ranging from 120 m to 1 km. At all scales, we found two coarseness gradients (PCA axes) based on the relative importance in the spectrum of large (> 50 km⁻¹), intermediate (30–50 km⁻¹), small (10–25 km⁻¹) and very small (<10 km⁻¹) spatial frequencies. Textural ordination based on Fourier spectra provides a powerful and consistent framework to identifying prominent scales of landscape patterns and to compare scaling properties across landscapes.     

Landscape connectivity and animal behavior: functional grain as a key determinant for dispersal

Landscape connectivity can be viewed from two perspectives that could be considered as extremes of a gradient: functional connectivity (refers to how the behavior of a dispersing organism is affected by landscape structure and elements) and structural connectivity (depends on the spatial configuration of habitat patches in the landscape like vicinity or presence of barriers). Here we argue that dispersal behavior changes with landscape configuration stressing the evolutionary dimension that has often been ignored in landscape ecology. Our working hypothesis is that the functional grain of resource patches in the landscape is a crucial factor shaping individual movements, and therefore influencing landscape connectivity. Such changes are likely to occur on the short-term (some generations). We review empirical studies comparing dispersal behavior in landscapes differing in their fragmentation level, i.e., with variable resource grain. We show that behavioral variation affecting each of the three stages of the dispersal process (emigration, displacement or transfer in the matrix, and immigration) is indeed likely to occur according to selective pressures resulting from changes in the grain of the landscape (mortality or deferred costs). Accordingly, landscape connectivity results from the interaction between the dispersal behavior of individuals and the grain of each particular landscape. The existence of this interaction requires that connectivity estimates (being based on individual-based models, least cost distance algorithms, and structural connectivity metrics or even Euclidian distance) should be carefully evaluated for their applicability with respect to the required level of precision in species-specific and landscape information.     

Multi-scale site and landscape effects on the vulnerable superb parrot of south-eastern Australia during the breeding season

The threatened superb parrot of south-eastern Australia exemplifies many of the challenges associated with research on wide-raging organisms which live ‘off-reserve’. Challenges include that most land is privately owned and that landscape use by such organisms does not always conform to traditional schematic and categorical landscape/fragmentation models. A multi-scale approach for embedding the detection of site-level and landscape context effects into landscape sampling design and subsequent statistical analysis is presented. The superb parrot was found scattered at varying densities throughout the agricultural landscapes of the South-West Slopes, much of which was privately owned. It responded to site-level variables and the surrounding landscape context. Overall, the superb parrot favoured lower elevation sites which were dominated by scattered, open woodlands, where Blakely’s red gum was a significant component. Mean plant productivity within 2 km, levels of woody tree cover within 3 km and (with caveats) length of roads within 3 km had a major effect on site-level response, indicating conditions in the surrounding local landscape are important to the superb parrot. This multi-scale response requires a multi-scale conservation and restoration strategy. The importance of open tree cover and amounts of Blakely’s red gum are a matter for concern, due to a general lack of tree regeneration and the particular susceptibility of Blakely’s red gum to dieback. The scattered trees in the agricultural matrix were important to the superb parrot, suggesting that it views these landscapes as a continuum of usable habitat. Strategies for restoration of larger habitat remnants should also include regeneration of trees in scattered pattern in the wider landscape, and Blakely’s red gum should be part of any strategy along with other key species such as yellow and white box. The landscape sampling approach successfully addressed the challenges of whole-landscape research. This highlights the value of ‘off-reserve’ studies across whole landscapes.     

Landscape effects on scales of movement by white-tailed deer in an agricultural–forest matrix

Understanding how organisms respond to landscape heterogeneity is foundational to landscape ecology. We characterized seasonal scales of movement of white-tailed deer (Odocoileus viginianus) in an agricultural–forest matrix using first-passage time analysis (FPT) for 62 GPS-collared individuals. We investigated whether those scales were driven by demographic or landscape features. We found FPT for each individual across all seasons was typically dominated by a peak in variance of FPT/area at scales (radii) from 425 to 1,675 m. These peaks occurred at scales consistent with seasonal space use. We observed additional lower magnitude peaks at larger scales (3,000–6,000 m) and small scales (25–150 m). Peaks at larger scales were associated with seasonal migrations and dispersal events. Small scale peaks may represent resting or foraging behavior. Female movements were organized at smaller scales than males in the spring/summer season. Models relating landscape features to movement scales suggest that deer perceive and move within the landscape differently as the roles of dominant land-cover types shift seasonally. During winter, configuration (interspersion/juxtaposition) of land-cover types is more important to deer than during spring/summer and fall. During spring/summer and fall, movement behavior may be dictated by reproductive and harvest activities.     

Mapping hotspots of multiple landscape functions: a case study on farmland afforestation in Scotland

Many conservation and restoration efforts in developed countries are increasingly based on the premise of recognising and stimulating more ‘multi-functionality’ in agricultural landscapes. Public policy making is often a pragmatic process that involves efforts to negotiate trade-offs between the potentially conflicting demands of various stakeholders. Conservationists’ efforts to influence policy making, can therefore benefit from any tool that will help them to identify other socio-economic functions or values that coincide with good ecological conservation options. Various types of socio-economic objectives have in recent years been mapped across landscapes and so there are now important opportunities to explore the spatial heterogeneity of these diverse functions across the wider landscape in search of potential spatial synergies, i.e. ‘multiple win locations’ or multifunctional ‘hotspots’. This paper explores the potential occurrence of such synergies within the agricultural landscape of northeast Scotland and evaluates an existing woodland planting policy using and combining three different policy objectives. Our results show that there are indeed broad areas of the studied landscape where multiple objectives (biodiversity, visual amenity and on-site recreation potential) could be achieved simultaneously (hotspots), and that the case study which we evaluate (the Farm Woodland Premium Scheme) could be much better spatially targeted with regards to each individual objective as well as with regards to these hotspots of multifunctionality.