Landscape attributes and life history variability shape genetic structure of trout populations in a stream network

Spatial and temporal landscape patterns have long been recognized to influence biological processes, but these processes often operate at scales that are difficult to study by conventional means. Inferences from genetic markers can overcome some of these limitations. We used a landscape genetics approach to test hypotheses concerning landscape processes influencing the demography of Lahontan cutthroat trout in a complex stream network in the Great Basin desert of the western US. Predictions were tested with population- and individual-based analyses of microsatellite DNA variation, reflecting patterns of dispersal, population stability, and local effective population sizes. Complementary genetic inferences suggested samples from migratory corridors housed a mixture of fish from tributaries, as predicted based on assumed migratory life histories in those habitats. Also as predicted, populations presumed to have greater proportions of migratory fish or from physically connected, large, or high quality habitats had higher genetic variability and reduced genetic differentiation from other populations. Populations thought to contain largely non-migratory individuals generally showed the opposite pattern, suggesting behavioral isolation. Estimated effective sizes were small, and we identified significant and severe genetic bottlenecks in several populations that were isolated, recently founded, or that inhabit streams that desiccate frequently. Overall, this work suggested that Lahontan cutthroat trout populations in stream networks are affected by a combination of landscape and metapopulation processes. Results also demonstrated that genetic patterns can reveal unexpected processes, even within a system that is well studied from a conventional ecological perspective.     

A comparison of metrics predicting landscape connectivity for a highly interactive species along an urban gradient in Colorado, USA

Many organisms persist in fragmented habitat where movement between patches is essential for long-term demographic and genetic stability. In the absence of direct observation of movement, connectivity or isolation metrics are useful to characterize potential patch-level connectivity. However, multiple metrics exist at varying levels of complexity, and empirical data on species distribution are rarely used to compare performance of metrics. We compared 12 connectivity metrics of varying degrees of complexity to determine which metric best predicts the distribution of prairie dog colonies along an urban gradient of 385 isolated habitat patches in Denver, Colorado, USA. We found that a modified version of the incidence function model including area-weighting of patches and a cost-weighted distance surface best predicted occupancy, where we assumed roads were fairly impermeable to movement, and low-lying drainages provided dispersal corridors. We also found this result to be robust to a range of cost weight parameters. Our results suggest that metrics should incorporate both patch area and the composition of the surrounding matrix. These results provide guidance for improved landscape habitat modeling in fragmented landscapes and can help identify target habitat for conservation and management of prairie dogs in urban systems.     

Identification of functional corridors with movement characteristics of brown bears on the Kenai Peninsula,Alaska

We identified primary habitat and functional corridors across a landscape using Global Positioning System (GPS) collar locations of brown bears (Ursus arctos). After deriving density, speed, and angular deviation of movement, we classified landscape function for a group of animals with a cluster analysis. We described areas with high amounts of sinuous movement as primary habitat patches and areas with high amounts of very directional, fast movement as highly functional bear corridors. The time between bear locations and scale of analysis influenced the number and size of corridors identified. Bear locations should be collected at intervals ≤6 h to correctly identify travel corridors. Our corridor identification technique will help managers move beyond the theoretical discussion of corridors and linkage zones to active management of landscape features that will preserve connectivity.     

Effects of area and isolation of woodland patches on herbaceous plant species richness across Great Britain

Richness of Ancient Woodland Indicator plant species was analysed in 308 woodland patches that were surveyed during the Countryside Survey of Great Britain carried out in 1998. The Countryside Survey recorded vegetation plots and landscape structure in 569 stratified 1 km sample squares and developed a remotely-sensed land cover map of the UK. Using these datasets, we tested the hypothesis that Ancient Woodland Indicator species richness in woodland fragments was limited by patch area, shape and spatial isolation and that woodland patches located in the lowland region of Great Britain would respond differently than those in the upland region. The variation in Ancient Woodland Indicator species richness in the British lowlands (n = 218) was mainly explained by patch area and two measures of connectivity, the length of hedgerows and lines of trees in the 1 km square and the area of woodland within 500 m of the vegetation plot. By contrast, variation in Ancient Woodland Indicator species richness in the British uplands (n = 90) was related to Ellenberg scores of the vegetation communities sampled – a surrogate for habitat quality – and no significant effect of spatial structure was detected. It therefore appears that the degree of fragmentation of woodland in the British lowlands limits the distribution of Ancient Woodland Indicator species, while in the uplands, failed colonisation is a matter of habitat quality rather than a result of landscape structure.This revised version was published online in May 2005 with corrections to the Cover Date.     

Effect of matrix habitat on the spread of flea beetle introductions for biological control of leafy spurge

Matrix habitats are known to influence the movement patterns of a variety of species but it is less well known whether these effects have strong implications for spatial population dynamics, including the spread of biological introductions. Using a spatially explicit simulation model parameterized with empirical data, we examine how grass and shrub matrix habitats, each offering different resistance to dispersal, influence the spread and impact of a biocontrol agent, Aphthona lacertosa, on the invasive weed, leafy spurge. Model predictions indicate that differential responses to matrix habitat have little effect on the agent’s spread over the study landscape and this is supported by statistical models fit to observed A. lacertosa incidence on the same landscape. Subsequent experimentation with the simulation model suggested that A. lacertosa colonization rates were largely unaffected by increases in amount of the more restrictive shrub matrix. However, simulations of an hypothetical species with greater overall dispersal ability but reduced dispersal rate through shrub matrix showed that colonization rates were noticeably reduced when the percentage of shrub matrix on the landscape approached 50%. Combined these results suggest that some tailoring of release strategies may be required to accommodate the unique dispersal capabilities of different biocontrol agents on particular release landscapes, but for A. lacertosa there appears to be little effect of matrix habitat structure on rates of spread.     

How far do birds disperse seeds in the degraded tropical landscape of Hong Kong,China?

Information on seed dispersal distances is critical for understanding plant species persistence in habitat fragments and vegetation recovery when disturbance is reduced. In the degraded upland landscape of Hong Kong (22°N), the bulbuls Pycnonotus sinensis and P. jocosus are responsible for a large proportion of seed movements. Dispersal distances were estimated from gut passage times (GPTs) and movement patterns determined by radio-telemetry. Estimates were also made for the hwamei, Garrulax canorus. Seven adult P. sinensis and four adult G. canorus were tracked in winter, and six juvenile P. sinensis, three juvenile P. jocosus and two juvenile G. canorus in summer. GPTs were 5–122 min in the bulbuls and 18–61 min in the hwamei. Most 10-min movements were <100 m for the bulbuls and <50 m for the hwamei, but the largest were >1300 m for both bulbuls and 940 m for the hwamei. Displacement-time graphs generally levelled off rapidly, with median displacements <60 m after an hour, except with summer juvenile bulbuls. Median displacements during median gut passage times for seeds from single-seeded fruits were 40, 116 and 131 m, respectively, for winter adult P. sinensis and summer juvenile P. sinensis and P. jocosus. Maximum observed displacements during the maximum measured GPTs were >1 km for all bulbuls. Estimated dispersal distances were shorter for hwameis. The radio-telemetry results were supplemented by 49 h of visual observations, during which 1,510 bird movements across open areas were observed, 64% by P. sinensis, 13% by P. jocosus, and 0.5% by G. canorus. The bulbuls, therefore, connect habitat fragments in upland Hong Kong for plants with fruits within their maximum gape width.     

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.     

Effect of landscape structure on anopheline mosquito density and diversity in northern Thailand: Implications for malaria transmission and control

The influence of landscape structure on anopheline mosquito density and diversity was studied in a comparison of agricultural and forested landscapes in northern Thailand. Agriculture locations had significantly higher landscape diversity, more patches, smaller mean patch sizes, and more complex patch shapes than forest locations. Mosquito collections were undertaken during both dry and wet seasons from October 1997 to December 1999. The density of two forest-associated species, Anopheles maculatus s.s. and Anopheles minimus s.l., both primary malaria vectors in Thailand, was significantly higher in forest locations in at least one season. The density of two paddy field-associated species, Anopheles aconitus and Anopheles hyrcanus group did not differ between locations. Anopheles aconitus is a secondary malaria vector and An. hyrcanus group is not considered as a vector in Thailand. The density of An. minimus s.l. was positively related to forest mean patch size, various water and paddy field landscape metrics and negatively related to landscape diversity. Anopheles hyrcanus group was also positively related to water metrics. Anopheline species diversity was negatively related to landscape diversity. Forest fragmentation resulting from human economic activities often increases landscape heterogeneity, which may result in a reduction in anopheline species diversity, as was the case in this study. There are indications that the effect of fruit orchards on anopheline diversity might be different in the dry season compared to the wet season. Fruit orchard landscape metrics affected species diversity negatively in the dry season and positively in the wet season. One reason for this could be that pesticides are typically applied in fruit orchards during the dry season. The conversion of forests to fruit orchards is a major land-use change in northern Thailand. These results show the complexity of vector status in northern Thailand and that vector and agriculture pest control are intricately interrelated. It is therefore important to include both the public health and agricultural sectors in controlling malaria vectors in the country. Our results also indicate that if landscape management should be used for malaria control in northern Thailand large-scale reduction and fragmentation of forest cover would be needed. Such drastic actions do not agree well with current global objectives concerning forest and biodiversity conservation This revised version was published online in July 2006 with corrections to the Cover Date.     

The Influence of Landscape Structure on Female Roe Deer Home-range Size

Animal distribution and abundance are greatly affected by the availability of their food resources, which also depends on landscape structure. Lothar hurricane in 1999 had profoundly modified the structure of the forests in France, affecting the habitat quality of ungulates. We tested whether the variations in home-range size of 23 female roe deer were influenced by the fragmentation of the landscape caused by Lothar in the Chizé forest, namely by the increase in heterogeneity associated with the localized massive tree felling. Home-range size was studied in the summers of 2001 and 2002 and we found that variation in home-range size was mainly explained by only one landscape variable: edge density. Home-range size decreased as edge density increased, which is consistent with the fact that edges are good browsing habitats for roe deer. The result of this study suggests that, after 2 years, the hurricane had improved the quality of the home ranges by creating more forest heterogeneity and increasing the contacts between the different vegetation patches within the home range. These results highlight the fact that spatial heterogeneity is likely to be a key factor influencing the distribution and local population density.     

Connectivity change in habitat networks

Habitat management is essential for safeguarding important flora and fauna. Further, habitat connectivity is a crucial component for maintaining biodiversity given that it is known to have implications for species persistence. However, damage to habitat due to natural and human induced hazards can alter spatial relationships between habitats, potentially impacting biodiversity. Therefore, the susceptibility of spatial relationships to patch loss and associated connectivity degradation is obviously an important factor in maintaining existing or planned habitat networks. Identifying patches vital to connectivity is critical both for effectively prioritizing protection (e.g., enhancing habitat connectivity) and establishing disaster mitigation measures (e.g., stemming the spread of habitat loss). This paper presents a methodology for characterizing connectivity associated with habitat networks. Methods for evaluating habitat network connectivity change are formalized. Examples are presented to facilitate analysis of connectivity in the management of biodiversity.

The influence of habitat availability and landscape structure on the distribution of wood cricket (Nemobius sylvestris) on the Isle of Wight, UK

Little information is available regarding the landscape ecology of woodland invertebrate species with limited dispersal ability. An investigation was therefore conducted within woodland fragments in an agricultural landscape for the flightless wood cricket (Nemobius sylvestris) on the Isle of Wight, UK. The current pattern of distribution of the species, established during a field survey, was related to measures of habitat availability and habitat isolation/fragmentation. Results revealed that wood cricket populations were patchily distributed and mainly found in relatively large mature woodland fragments situated closely (<50 m) to another occupied site. Although the occurrence of wood cricket was related to fragment area, isolation, habitat availability and woodland age, a logistic regression model revealed that presence of the species was most accurately predicted by fragment isolation and area alone. These results highlight the vulnerability of relatively immobile woodland invertebrate species, such as wood cricket, to the impacts of habitat loss and fragmentation.     

Distribution of the Cinnabar moth Tyria jacobaeae L. at landscape scale: use of linear landscape structures in egg laying on larval hostplant exposures

The distribution of the xerothermophilous Cinnabar moth Tyria jacobaeae was studied in a low mountain region in western Germany between 1989 and 2001. T. jacobaeae started its immigration into the study area in 1989 and first established populations in climatically favoured habitats like abandoned quarries and train stations where the larval host plant, ragwort (Senecio jacobaea), occurs. Analysis of landscape features (altitude, morphology) reveals that T. jacobaeae then dispersed along valleys with roads to higher altitudes of the study area. Elevations of occupied sites increased between 1989 and 2001. In order to investigate whether dispersal is affected by linear structures like valleys or roads with gravelled verges, hostplant exposures were placed at a distance of 600 m to the next population of T. jacobaeae. The experiments suggest that egglaying predominantly took place on exposures in valleys with roads and sparsely plant-covered verges but can also occur along valleys lacking roads and suitable habitats. However, larvae were never recorded on Senecio exposures which were placed aside from valleys and roads.This revised version was published online in May 2005 with corrections to the Cover Date.     

Effects of corridors on home range sizes and interpatch movements of three small mammal species

Corridors are predicted to benefit populations in patchy habitats bypromoting movement, which should increase population densities, gene flow, andrecolonization of extinct patch populations. However, few investigators haveconsidered use of the total landscape, particularly the possibility ofinterpatch movement through matrix habitat, by small mammals. This studycompares home range sizes of 3 species of small mammals, the cotton mouse(Peromyscus gossypinus), old-field mouse (P.polionotus) and cotton rat (Sigmodon hispidus)between patches with and without corridors. The study site was in S. Carolina,USA. Corridor presence did not have astatistically significant influence on average home range size. Habitatspecialization and sex influenced the probability of an individual movingbetween 2 patches without corridors. The results of this study suggest thatsmall mammals may be more capable of interpatch movement without corridors thanis frequently assumed.This revised version was published online in May 2005 with corrections to the Cover Date.     

Evaluating effects of low quality habitats on regional population growth in <Emphasis Type="Italic">Peromyscus leucopus</Emphasis>: Insights from field-parameterized spatial matrix models

Due to complex population dynamics and source–sink metapopulation processes, animal fitness sometimes varies across landscapes in ways that cannot be deduced from simple density patterns. In this study, we examine spatial patterns in fitness using a combination of intensive field-based analyses of demography and migration and spatial matrix models of white-footed mouse (Peromyscus leucopus) population dynamics. We interpret asymptotic population growth rates from these spatial models as fitness-based measures of habitat-quality and use elasticity analysis to further explore model behavior and the roles of migration. In addition, we compare population growth rates at the spatial scale of single habitats and the landscape-level scale at which these habitats are assembled. To this end, we employ emerging techniques in multi-scale estimation of demography and movement and recently described vec-permutation methods for spatial matrix notation and analysis. Our findings indicate that the loss of low quality habitats or reductions in movement from these habitats into higher quality areas could negatively affect landscape-level population fitness.     

Importance of Wetland Landscape Structure to Shorebirds Wintering in an Agricultural Valley

Only recently has the influence of landscape structure on habitat use been a research focus in wetland systems. During non-breeding periods when food can be locally limited, wetland spatial pattern across a landscape may be of great importance in determining wetland use. We studied the influence of landscape structure on abundances of wintering Dunlin (Calidris alpina) and Killdeer (Charadrius vociferus) observed on wetlands in the agricultural Willamette Valley of Oregon, USA, during two winters (1999–2000, 2000–2001) of differing rainfall. We examined (1) shorebird use within a sample of 100 km² regions differing in landscape structure (hectares of shorebird habitat [wet, unvegetated]) and (2) use of sites differing in landscape context (area of shorebird habitat within a species-defined radius). For use of sites, we also assessed the influence of two local characteristics: percent of soil exposed and area of wet habitat. We analyzed data using linear regression and information-theoretic modeling. During the dry winter (2000–2001), Dunlin were attracted to regions with more wetland habitat and their abundances at sites increased with greater area of shorebird habitat within both the site and the surrounding landscape. In contrast, Dunlin abundances at sites were related to availability of habitat at only a local scale during the wet winter (1999–2000). Regional habitat availability was of little importance in predicting Killdeer distributions, and Killdeer site use appeared unrelated to habitat distributions at both landscape and local scales. Results suggest prioritizing sites for conservation that are located in areas with high wetland coverage.     

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

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

Are hedgerows the route to increased farmland small mammal density? Use of hedgerows in British pastoral habitats

Linear habitats are becoming increasingly common as a consequence of habitat fragmentation, and may provide the sole habitat for some species. Hedgerows are linear features that can vary substantially in structure and quality. Having surveyed 180 hedgerows, in four locations, and sampled their small mammal communities we examined the effect of physical hedgerow attributes on the abundance of small mammal species. Using three elements of landscape structure, we explored whether variation was best explained by the Random Sample Hypothesis (that small islands represent a random sample of those species populating larger areas), or by the Fragmentation Hypothesis (that species abundance will decrease with a loss of habitat area). We tested the relationship between the relative abundance of small mammals and 1. hedgerow connectivity; 2. total habitat availability and 3. local habitat complexity. We then explored the predictive power of combinations of these habitat variables. Connectivity was a positive predictor of wood mice Apodemus sylvaticus, and hedgerow gappiness was a negative predictor of bank voles Clethrionomys glareolus. The total amount of habitat available (hedgerow width, height and length) was a positive indicator of total small mammal biomass. These results support the Fragmentation Hypothesis that species abundance and distribution decrease with a loss of habitat area. The preservation of linear and associated habitats may therefore be important in maintaining metapopulations of the species we studied.     

Use of space by the yellow-footed antechinus, Antechinus flavipes, in a fragmented landscape in South Australia

An understanding of how individual species are able to persist and move within fragmented landscapes is critical for elucidating the effects of fragmentation and aiding in the management of species. Here, we studied movement behaviour of the dasyurid Antechinus flavipes in a heavily fragmented landscape using trapping and radiotracking. We assessed the ability of animals to move within and amongst small (<6 ha) remnants and make use of the matrix, and investigated how females used the available space within remnants. Seventeen between-remnant movements were detected from 428 recaptures, ranging in length from 30 to 720 m and averaging 352 m. Most were by adult males during the breeding season, with 40% more than 500 m. Landscape types traversed would have included exotic pine plantations, open grazed areas and roads. Between-site movements of juveniles were only detected on three occasions. However, few young males were captured as adults, suggesting high dispersal rates and considerable matrix use. Conversely, despite high female recapture rates, again only three between-site movements were recorded. Radiotracking further indicated that females confined foraging to remnants, with occasional forays to isolated trees in paddocks. Female home range areas were similar for remnants and forest (0.04–0.66 ha). A. flavipes is clearly able to persist in very small patches of native vegetation in the landscape studied here. Its long-term persistence appears dependent on the ability of females to maintain a presence in the small remnants, and of unrelated males to move between remnants to breed with resident females. This study illustrates the importance of recognising the occurrence of metapopulations in fragmented landscapes for conservation management.     

Is bird incidence in Atlantic forest fragments influenced by landscape patterns at multiple scales?

The degree to which habitat fragmentation affects bird incidence is species specific and may depend on varying spatial scales. Selecting the correct scale of measurement is essential to appropriately assess the effects of habitat fragmentation on bird occurrence. Our objective was to determine which spatial scale of landscape measurement best describes the incidence of three bird species (Pyriglena leucoptera, Xiphorhynchus fuscus and Chiroxiphia caudata) in the fragmented Brazilian Atlantic forest and test if multi-scalar models perform better than single-scalar ones. Bird incidence was assessed in 80 forest fragments. The surrounding landscape structure was described with four indices measured at four spatial scales (400-, 600-, 800- and 1,000-m buffers around the sample points). The explanatory power of each scale in predicting bird incidence was assessed using logistic regression, bootstrapped with 1,000 repetitions. The best results varied between species (1,000-m radius for P. leucoptera; 800-m for X. fuscus and 600-m for C. caudata), probably due to their distinct feeding habits and foraging strategies. Multi-scale models always resulted in better predictions than single-scale models, suggesting that different aspects of the landscape structure are related to different ecological processes influencing bird incidence. In particular, our results suggest that local extinction and (re)colonisation processes might simultaneously act at different scales. Thus, single-scale models may not be good enough to properly describe complex pattern–process relationships. Selecting variables at multiple ecologically relevant scales is a reasonable procedure to optimise the accuracy of species incidence models.