Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.     

Fine‐scale habitat selection by sympatric Canada lynx and bobcat

The Canada lynx (Lynx canadensis) and the bobcat (Lynx rufus) are closely related species with overlap at their range peripheries, but the factors that limit each species and the interactions between them are not well understood. Habitat selection is a hierarchical process, in which selection at higher orders (geographic range, home range) may constrain selection at lower orders (within the home range). Habitat selection at a very fine scale within the home range has been less studied for both lynx and bobcat compared to selection at broader spatiotemporal scales. To compare this fourth‐order habitat selection by the two species in an area of sympatry, we tracked lynx and bobcat during the winters of 2017 and 2018 on the north shore of Lake Huron, Ontario. We found that both lynx and bobcat selected shallower snow, higher snowshoe hare abundance, and higher amounts of coniferous forest at the fourth order. However, the two species were spatially segregated at the second order, and lynx were found in areas with deeper snow, more snowshoe hare, and more coniferous forest. Taken together, our findings demonstrate that the lynx and bobcat select different resources at the second order, assorting along an environmental gradient in the study area, and that competition is unlikely to be occurring between the two species at finer scales.     

Habitat selection and the value of information in heterogenous landscapes

Despite the wide usage of the term information in evolutionary ecology, there is no general treatise between fitness (i.e. density‐dependent population growth) and selection of the environment sensu lato. Here we 1) initiate the building of a quantitative framework with which to examine the relationship between information use in spatially heterogeneous landscapes and density‐dependent population growth, and 2) illustrate its utility by applying the framework to an existing model of breeding habitat selection. We begin by linking information, as a process of narrowing choice, to population growth/fitness. Second, we define a measure of a population's penalty of ignorance based on the Kullback–Leibler index that combines the contributions of resource selection (i.e. biased use of breeding sites) and density‐dependent depletion. Third, we quantify the extent to which environmental heterogeneity (i.e. mean and variance within a landscape) constrains sustainable population growth of unbiased agents. We call this the heterogeneity‐based fitness deficit, and combine this with population simulations to quantify the independent contribution of information‐use strategies to the total population growth rate. We further capitalize on this example to highlight the interactive effects of information between ecological scales when fear affects individual fitness through phenotypic plasticity. Informed breeding habitat selection moderates the demographic cost of fear commensurate with density‐dependent information use. Thus, future work should attempt to differentiate between phenotypic plasticity (i.e. acute fear) and demographic responses (i.e. chronic changes in population size). We conclude with a broader discussion of information in alternative contexts, and explore some evolutionary considerations for information use. We note how competition among individuals may constrain the information state among individuals, and the implications of this constraint under environmental change.     

Neutral communities in fragmented landscapes

Habitat fragmentation displays a crucial role in conservation biology. Despite this, little is known about the detailed ecological consequences of habitat fragmentation due to the scarce number of controlled experimental surveys. The species–area relationship, a fundamental concept in ecology, requires the understanding of the fragmentation effects in a long term perspective, which turns this task even harder. Here we address the spatial patterns of species distribution in fragmented landscapes, assuming a neutral community model. We study the species area relationship and how its shape changes as the landscape becomes more fragmented. Recent investigations, based on extensive computer simulation, have contributed to establish some definite conclusions in the study of non‐fragmented landscapes: the existence of a three‐regime or two‐regime scenario for the species–area relationship, the emergence of a power‐law regime at intermediate scales and the augment of the species–area exponent z with the speciation rate. Despite the recent efforts, some other questions remain, such as the dependence of z in the whole range of the speciation rate. Questions like these are currently debated but generalizations cannot be drawn. This is the first paper, to our knowledge, that uses the coalescence method and neutral theory to examine biodiversity on more complex spatial structures. Our simulation results corroborate that the fragmentation plays a crucial role in shaping the species–area relationship, by determining the existence and extension of the power‐law regime associated with small and intermediate areas. On the other hand, when individuals are allowed to disperse over longer distances the species–area relationship now displays the classic triphasic pattern, and the intermediate regime, which is well described by a power‐law, is established even for highly fragmented landscapes.     

Habitat selection by Eurasian lynx (Lynx lynx) is primarily driven by avoidance of human activity during day and prey availability during night

The greatest threat to the protected Eurasian lynx (Lynx lynx) in Central Europe is human‐induced mortality. As the availability of lynx prey often peaks in human‐modified areas, lynx have to balance successful prey hunting with the risk of encounters with humans. We hypothesized that lynx minimize this risk by adjusting habitat choices to the phases of the day and over seasons. We predicted that (1) due to avoidance of human‐dominated areas during daytime, lynx range use is higher at nighttime, that (2) prey availability drives lynx habitat selection at night, whereas high cover, terrain inaccessibility, and distance to human infrastructure drive habitat selection during the day, and that (3) habitat selection also differs between seasons, with altitude being a dominant factor in winter. To test these hypotheses, we analyzed telemetry data (GPS, VHF) of 10 lynx in the Bohemian Forest Ecosystem (Germany, Czech Republic) between 2005 and 2013 using generalized additive mixed models and considering various predictor variables. Night ranges exceeded day ranges by more than 10%. At night, lynx selected open habitats, such as meadows, which are associated with high ungulate abundance. By contrast, during the day, lynx selected habitats offering dense understorey cover and rugged terrain away from human infrastructure. In summer, land‐cover type greatly shaped lynx habitats, whereas in winter, lynx selected lower altitudes. We concluded that open habitats need to be considered for more realistic habitat models and contribute to future management and conservation (habitat suitability, carrying capacity) of Eurasian lynx in Central Europe.     

Habitat restoration will help some functional plant types persist under climate change in fragmented landscapes

In the next century, global climate change is predicted to have large influences on species' distributions. Much of the research in this area has focused on predicting the areas where conditions will be suitable for the species in future, and thus the potential distribution of the species. However, it is equally important to predict the relative abilities of species to migrate into new suitable areas as conditions shift, while accounting for dynamic processes, such as dispersal, maturation, mortality, and reproduction, as well as landscape characteristics, such as level of habitat fragmentation and connectivity. In this study, we developed a spatially explicit individual‐based model that addresses these factors. As a motivating case study, we based aspects of the model on southwest Australia, a global biodiversity hotspot, but stress that the results obtained are generalizable beyond this region. Using the model, we enhanced current understanding of climate change impacts by investigating how and to what extent the functional traits of plant species affect their ability to move with climate change across landscapes with various levels of fragmentation. We also tested the efficacy of strategic restoration, such as planting corridors to increase connectivity among fragments. We found that even if the landscape is fully intact, only an average of 34.2% of all simulated functional groups had a good chance of successfully tracking climate change. However, our study highlights the power of strategic restoration as a tool for increasing species persistence. Corridors linking fragments increased species persistence rates by up to 24%. The lowest persistence rates were found for trees, a functional group with high dispersal but also long generation times. Our results indicate that for trees intervention techniques, such as assisted migration might be required to prevent species losses.     

Seed-dispersal distributions by trumpeter hornbills in fragmented landscapes

Frugivorous birds provide important ecosystem services by transporting seeds of fleshy fruited plants. It has been assumed that seed-dispersal kernels generated by these animals are generally leptokurtic, resulting in little dispersal among habitat fragments. However, little is known about the seed-dispersal distribution generated by large frugivorous birds in fragmented landscapes. We investigated movement and seed-dispersal patterns of trumpeter hornbills (Bycanistes bucinator) in a fragmented landscape in South Africa. Novel GPS loggers provide high-quality location data without bias against recording long-distance movements. We found a very weakly bimodal seed-dispersal distribution with potential dispersal distances up to 14.5 km. Within forest, the seed-dispersal distribution was unimodal with an expected dispersal distance of 86 m. In the fragmented agricultural landscape, the distribution was strongly bimodal with peaks at 18 and 512 m. Our results demonstrate that seed-dispersal distributions differed when birds moved in different habitat types. Seed-dispersal distances in fragmented landscapes show that transport among habitat patches is more frequent than previously assumed, allowing plants to disperse among habitat patches and to track the changing climatic conditions.     

Patch-occupancy dynamics in fragmented landscapes

Recent work on the dynamics of species living In fragmented landscapes has produced much Information on patterns of habitat patch occupancy in a wide range of organisms. Building on an elementary Markov chain model of patch occupancy, a family of Incidence-function models can be constructed for particular kinds of metapopulations. These models can be parameterized with field data on patch occupancy, and the models can be used to make quantitative predictions about specific metapopulations. This approach provides a potentially powerful tool for the management of reserve networks and species living in fragmented landscapes.     

Forest-climate interactions in fragmented tropical landscapes

In the tropics, habitat fragmentation alters forest-climate interactions in diverse ways. On a local scale (less than 1 km), elevated desiccation and wind disturbance near fragment margins lead to sharply increased tree mortality, thus altering canopy-gap dynamics, plant community composition, biomass dynamics and carbon storage. Fragmented forests are also highly vulnerable to edge-related fires, especially in regions with periodic droughts or strong dry seasons. At landscape to regional scales (10-1000 km), habitat fragmentation may have complex effects on forest-climate interactions, with important consequences for atmospheric circulation, water cycling and precipitation. Positive feedbacks among deforestation, regional climate change and fire could pose a serious threat for some tropical forests, but the details of such interactions are poorly understood.     

Dispersal and extinction in fragmented landscapes

Evolutionary and population dynamics models suggest that the migration rate will affect the probability of survival in fragmented landscapes. Using data for butterfly species in the fragmented British landscape and in immediately adjoining areas of the European continent, this paper shows that species of intermediate mobility have declined most, followed by those of low mobility, whereas high-mobility species are generally surviving well. Compared to the more sedentary species, species of intermediate mobility require relatively large areas where they breed at slightly lower local densities. Intermediate mobility species have probably fared badly through a combination of metapopulation (extinction and colonization) dynamics and the mortality of migrating individuals which fail to find new habitats in fragmented landscapes. Habitat fragmentation is likely to result in the non-random extinction of populations and species characterized by different levels of dispersal, although the details are likely to depend on the taxa, habitats and regions considered.     

Habitat use by beech martens in a fragmented landscape

We investigated key habitat features used by beech martens occupying a fragmented agricultural landscape. Locations of eight martens were radio-monitored daily for 4–12 months each. We evaluated the selection of home ranges within a geographic area, as well as the selection of habitat features within home ranges, using random simulations for comparison. Home ranges included more wood and scrub vegetation, and less arable land, than random simulations. They also included a higher proportion of watercourses with continuous vegetation along their verges, and were closer to these watercourses than random simulations. Within home ranges, beech martens used wood and scrub vegetation, and farm buildings, more intensively than expected, and arable land less than expected. On average, beech martens stayed close to watercourses (where most of the wood vegetation occurred) but not to farm buildings, indicating that only the former shaped beech marten movements. By these findings, arable land appeared to act as a seldom traversed barrier, making farm buildings uneasy to reach if they are located far from watercourses.     

The matrix matters: effective isolation in fragmented landscapes

Traditional approaches to the study of fragmented landscapes invoke an island-ocean model and assume that the nonhabitat matrix surrounding remnant patches is uniform. Patch isolation, a crucial parameter to the predictions of island biogeography and metapopulation theories, is measured by distance alone. To test whether the type of interpatch matrix can contribute significantly to patch isolation, I conducted a mark-recapture study on a butterfly community inhabiting meadows in a naturally patchy landscape. I used maximum likelihood to estimate the relative resistances of the two major matrix types (willow thicket and conifer forest) to butterfly movement between meadow patches. For four of the six butterfly taxa (subfamilies or tribes) studied, conifer was 3-12 times more resistant than willow. For the two remaining taxa (the most vagile and least vagile in the community), resistance estimates for willow and conifer were not significantly different, indicating that responses to matrix differ even among closely related species. These results suggest that the surrounding matrix can significantly influence the "effective isolation" of habitat patches, rendering them more or less isolated than simple distance or classic models would indicate. Modification of the matrix may provide opportunities for reducing patch isolation and thus the extinction risk of populations in fragmented landscapes.     

Making statistics biologically relevant in fragmented landscapes

The biological impacts of habitat fragmentation are routinely assessed using standard statistical modelling techniques that are used across many ecological disciplines. However, to assess the biological relevance of fragmentation impacts, we must consider an extra, spatial dimension to the standard statistical model: the biological importance of a significant and well supported model with large effect sizes crucially depends on the configuration of habitat within the study area. We argue that mapping the outputs from statistical models across a study area generates biologically meaningful estimates of fragmentation impacts. Integrating traditional statistical approaches with geographic information systems will facilitate rigorous comparisons of fragmentation impacts between taxa, studies and ecosystems.     

Habitat selection by helminths: a hypothesis

How do liver worms find the liver? Why are heartworms always found in the heart? Attempts to answer these questions have invariably yielded inconclusive results. A major problem has been the use of models or hypotheses derived from studies of free-living organisms. The parasite's environment is distinct from free-living environments in several fundamental ways and these differences will impact on the evolution of parasite behaviours. In this article, Michael Sukhdeo outlines a hypothesis for habitat selection behavior of helminths that is based on the specific environmental conditions within the host.     

Black box and attics: Habitat selection and resource use by large threatened pythons in landscapes with contrasting human modification

Woodland and forest degradation, driven predominately by agricultural and pastoral production, is a crisis facing many species globally, in particular hollow‐dependent fauna. Large predatory species play important roles in both ecosystems and conservation strategies, but few studies have examined habitat selection of such species in intensively human‐modified landscapes. We quantified habitat selection and resource use by a large, top‐order and threatened snake (carpet python, Morelia spilota), between adjacent areas of high and low anthropogenic modification in inland Australia, a region that has undergone considerable alteration since European settlement. At the low‐impact site, snakes preferred tree hollows and a structurally complex understorey, whereas at the high‐impact site, snakes preferred homestead attics. Based on the decline of the species in this region, however, high‐impact landscapes may only support snakes when they are adjacent to low‐impact habitats. Invasive species comprised a large part of snake diets in both landscape types. Carpet pythons, with large home ranges and habitat requirements that overlap with many smaller threatened mammalian and avian fauna, are generally well liked and easily identifiable by rural landholders. Accordingly, they may play a key role in conservation strategies aimed at the protection of woodland and hollow‐dependent fauna in heavily modified landscapes of Australia's inland regions. However, invasive species, which tend to contribute to declines in native species inhabiting arid and semi‐arid Australia, are beneficial and important to pythons. Our study therefore highlights the diversity of effects that two major threats to biodiversity – habitat loss and invasive species – can have on different species within the same ecosystem.     

Density dependence slows invader spread in fragmented landscapes

Patchiness is a defining characteristic of most natural and anthropogenic habitats, yet much of our understanding of how invasions spread has come from models of spatially homogeneous environments. Except for populations with Allee effects, an invader's growth rate when rare and dispersal determine its spread velocity; intraspecific competition has little to no influence. How this result might change with landscape patchiness, however, is poorly understood. We used simulation models and their analytical approximations to explore the effect of density dependence on the spread of annual plant invaders moving through heterogeneous landscapes with gaps in suitable habitat. We found that landscape patchiness and discrete invader population size interacted to generate a strong role for density dependence. Intraspecific competition greatly slowed the spread of invasions through patchy landscapes by regulating how rapidly a population could produce enough seeds to surpass habitat gaps. Populations with continuously varying density showed no such effect of density dependence. We adapted a stochastic dispersal model to approximate spread when gap sizes were small relative to the mean dispersal distance and a Markov chain approximation for landscapes with large gaps. Our work suggests that ecologists must consider reproduction at both low and high densities when predicting invader spread.     

Sex-specific indicators of landscape use by servals: Consequences of living in fragmented landscapes

The conversion of wetlands and its associated habitat for farming, residential development and commercial purposes has led to many small disconnected patches of native vegetation surrounded by generally inhospitable human altered land uses. Such human interventions pose a serious threat to the survival of wetland dependent indicator species such as the serval (Leptailurus serval) in the Drakensberg Midlands of KwaZulu-Natal, South Africa. To address the effects of habitat fragmentation on serval space use we used GPS fixes of collared individuals for 100 days between May 2013 and January 2014. We tracked five females and eleven male servals with GPS-UHF collars. Fragmentation indices at the class level (wetland, forest with bushland, grassland and cropland) were measured within the collared serval's minimum convex polygon area (MCP) of home range. Serval use points and fragmentation indices were extracted to each sub-landscape unit of 2 km² falling within the MCP home range. The response of serval population to fragmentation indices was analyzed using generalized linear models at three levels: male, female and the overall population. We found that core area of wetland positively explained landscape use by servals. Effects of forest core area, forest proximity and patch richness were important for landscape use by male servals. Male and female serval use declined with increase in complex shapes of forest and grassland, wetland clumpiness for females and overall, and cropland patches for servals overall. Our results showed that wetland is a key determinant for the survival of male and female servals in fragmented landscapes therefore effective conservation of serval population demands adequate viable native habitat.     

Dispersal abilities and spatial patterns in fragmented landscapes

Recent theoretical studies suggest that the distribution of species in space has important implications for the conservation of communities in fragmented landscapes. Facilitation and dispersal are the primary mechanisms responsible for the formation of spatial patterns. Furthermore, disruptions in the formation of patterns arise after degradation, which can serve as an early indicator of stress in plant communities. Spatial dispersal ability and pattern formation were evaluated in 53 linear transects of 500 m in length within 14 fragments of natural vegetation within a matrix of abandoned crop fields in Cabo de Gata National Park, Almería, Spain. Fragments were classified into three size classes (< 300, 300–900, and > 900 ha). Fragment connectivity was quantified using the distances between fragments. Spatial dispersal ability was quantified for the 187 species recorded in the study. Species with restricted dispersal had the highest degree of long‐range spatial autocorrelation and, species that disperse by biotic vectors (e.g. vertebrates), the lowest. In addition, species most susceptible to fragmentation are vertebrate‐dispersed shrubs, which declined in abundance and was associated with loss of spatial organization in the smallest fragments. It is postulated that the positive feedback between abundance of recruitment and vertebrate visits influences the colonization and persistence of vertebrate‐dispersed shrubs, explaining its abundance in large fragments. Indeed, fragments lower than a certain threshold reduced spatial organization not only in shrubs with biotic dispersal, but also in species with abiotic dispersal (mainly wind) and with restricted dispersal. Fragments lower than a certain threshold may be vulnerable to a cascade of species loss because of reduced recruitment, establishment and patch biomass as a result of natural senescence, finally breaking up facilitative plant interactions. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 935–947.