Evaluating habitat connectivity methodologies: a case study with endangered African wild dogs in South Africa

Context

In fragmented landscapes, connectivity between subpopulations is vital for species’ persistence. Various techniques are used to assess the degree of connectivity between habitat patches, yet their performance is seldom evaluated. Models are regularly based on habitat selection by individuals in resident populations, yet dispersers may not require habitat which supports permanent residence.

Objectives and methods

Using a database of African wild dog (Lycaon pictus) occurrence records in north-eastern South Africa (n = 576), we developed and compared ecological niche models (ENM) for wild dogs packs and dispersers. Additionally, we used least cost path (LCP) and current flow models to assess connectivity. Results were further validated using occurrence records (n = 339) for cheetah (Acinonyx jubatus).

Results and conclusions

The ENM for wild dog packs identified large but isolated patches of suitable habitat, while the disperser ENM had greater suitability values for areas in between highly suitable patches. Without disperser-specific data, models omitted large areas which were confirmed to have provided connectivity. Although models derived from a potentially subjective cost layer have been criticised, the current flow model outperformed the other connectivity techniques and provided the most meaningful predictions for conservation planning. We identified five priority conservation areas for wild dogs, two of which had a greater feasibility for recolonisation. The scarcity of disperser-specific data promotes models using habitat data for resident individuals but here we illustrate the pitfalls thereof. Our study provides insights into the performance of these frequently employed techniques and how they may affect conservation management decisions.

     

Functional connectivity and matrix quality: network analysis for a critically endangered New Zealand lizard

Agricultural modification commonly leads to reductions in vegetation matrix quality and a resultant decrease in functional connectivity. In this study, a network analysis approach was used to assess the impact of agriculturally-induced reductions in vegetation matrix quality on the metapopulation dynamics of the critically endangered New Zealand grand skink (Oligosoma grande). Vegetation matrix quality was quantified in four sites exhibiting differing levels of modification within indigenous tussock grasslands in eastern Central Otago, New Zealand. Grand skink occupancy probability exhibited a positive correlation with the structural connectivity of primary habitat within the more modified study sites, whereas in the least modified site a complex matrix appeared to compensate for low structural connectivity. Results from this research indicate that the matrix is an important determinant of grand skink metapopulation dynamics and that an intricate balance exists between structural connectivity and the quality of the vegetation matrix. This research highlights the importance of assessing the impact of the matrix for individual species, particularly for conservation management.     

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.     

Towards a measure of functional connectivity: local synchrony matches small scale movements in a woodland edge butterfly

This study investigates the sensitivity of local synchrony to movement patterns of the Ringlet butterfly (Aphantopus hyperantus). We examine whether population synchrony, describing the correlated fluctuations of conspecific populations, may prove an effective surrogate measure for monitoring functional connectivity in this species without the requirement of exhaustive sampling. We compared the effect on population synchrony of two different distance measures, direct (Euclidean) distance and distance via woodland rides and edges, and also of habitat matrix composition. Population synchrony of A. hyperantus was calculated as the pairwise correlation between population time-series using 20?years of data from UK butterfly monitoring scheme transects. Local population synchrony was better explained by distance via woodland edges than direct distance, especially for woodland-dominated transects. These results are consistent with mark-recapture data previously collected on the Ringlet butterfly. The results indicate a sensitivity of population synchrony to butterfly local dispersal behaviour, particularly, to the use of habitat corridors and other functional dispersal routes. Population synchrony is considered to have potential as a surrogate measure of functional connectivity. With development, this method could become a valuable conservation tool for identifying important landscape features which promote species?? connectivity.     

Effects of landscape composition and connectivity on the distribution of an endangered parrot in agricultural landscapes

The extent and connectivity of individual habitat types strongly affects the distribution and abundance of organisms. However, little is known of how the level of connectivity and the interactions between different habitat types influences the distribution of species. Here, we used the geographically restricted and endangered regent parrot Polytelis anthopeplus monarchoides as a case study to examine the importance of composition and connectivity between different elements in 39 complex landscape mosaics (each 10 km radius). We compiled a database of 674 regent parrot nesting records, regional vegetation maps and measures of multipath connectivity between core vegetation types under different scenarios of resistance to movement provided by landscape elements. The occurrence of regent parrot nests was strongly affected by landscape composition, being positively related to the extent Eucalyptus camaldulensis riverine forest, but negatively related to the extent of semi-arid woodlands dominated by Eucalyptus largiflorens. Connectivity between E. camaldulensis forest (principal nesting habitat) and mallee (preferred feeding habitat) was a strong predictor of nest locations. Our study shows that the suitability of fragmented agricultural landscapes for supporting species can be greatly affected by connectivity and interactions between preferred and non-preferred habitats. For species that require complementary habitats such as the regent parrot, conservation management activities may be ineffective if they simply focus on a single core habitat type or the impacts of human land uses without regard to the interrelationships among landscape elements. While increasing the amount of primary preferred habitat should remain a cornerstone goal, increasing the extent and improving connectivity with alternative landscape elements also should be priority management objectives.     

Altered functional connectivity and genetic diversity of a threatened salamander in an agroecosystem

Context

Amphibian metapopulations have become increasingly fragmented in the Midwestern United States, with wetland-breeding salamanders being especially dependent on intact, high-quality forested landscapes. However, the degree to which amphibian populations are isolated, the factors that influence dispersal and, ultimately, functional connectivity remain areas in need of investigation.

Objectives and methods

We combined population demographic and genetic approaches to assess how a landscape fragmented by agriculture influences functional connectivity and metapopulation dynamics of a locally threatened salamander (Ambystoma jeffersonianum).

Results

We found that the allelic richness and heterozygosity of this species was significantly related to the level of connectivity with other occupied breeding wetlands and that decreased connectivity resulted in increased genetic differentiation. We also found that effective population size appears to be declining and, while correlative, our focal landscape has experienced significant losses of forested upland habitats and potential wetland breeding habitats over the last 200 years.

Conclusions

By combining population and landscape genetic analyses with an assessment of regional wetland occupancy, our study has uniquely synthesized genetic and metapopulation processes, while also incorporating the effects of the landscape matrix on dispersal, connectivity, and population differentiation. The significant relationship between connectivity with heterozygosity, allelic richness, and genetic divergence observed in this study reinforces empirical observations of long distance dispersal and movements in ambystomatid salamanders. However, our results show that protection of core habitat around isolated wetlands may not sufficiently minimize genetic differentiation among populations and preserve critical genetic diversity that may be essential for the long-term persistence of local populations.

     

Drones for butterfly conservation: larval habitat assessment with an unmanned aerial vehicle

Context

Evidence-based nature conservation focuses on ecological facts and the incorporation of knowledge on the ecology of species, including its entire life cycle. In butterflies, imagos and its larvae often demand specific and diverging micro-habitat structures and resources. In consequence, ecological requirements of the imaginal and pre-imaginal stage have to be taken into consideration to conduct effective conservation management.

Objective

Here we analyse ecological pre-requisites of imagos and larvae for two lycaenid butterfly species, the common blue Polyommatus icarus and the adonis blue Polyommatus bellargus. Both butterfly species occur in calcareous grasslands and mainly depend on two plant species at our study site, the horseshoe vetch Hippocrepis comosa and bird’s-foot trefoil Lotus corniculatus. These plant species serve as nectar sources and larval host plants for the two butterfly species.

Methods

First, we assessed the occurrence of imagines and larvae of the two butterfly species and recorded various micro-habitat characteristics, like the number of flower buds of the two main host plants, the surrounding vegetation height, percentage of bare soil, availability of shadow, and the distance to and geographic direction of thickets at respective sites. In a second step we took high resolution aerial pictures from our study area using an unmanned aerial vehicle (drone). Based on these aerial pictures and the information on the larvae´s habitat preference from our field observations, we trained a habitat suitability model to identify micro-habitat structures suitable for larvae of the two butterfly species.

Results

We found that abundance of imagos is positively correlated with flower bud density of the two host plants. Low vegetation height and high proportion of bare soil (but not flower bud density) positively influence egg oviposition. The calculated habitat suitability models predict the occurrence of high quality larval habitats with high prediction power (AUC = 0.72).

Conclusions

This combined data set consisting of field observations, high resolution aerial pictures taken from an unmanned aerial vehicle, and models underline that (1) species with complex life cycles may request more than one habitat niche, depending its stage of development, and (2) high resolution aerial pictures taken from drones provide valuable background data to generate habitat suitability models—even on a micro scale but covering larger parts of a landscape.

     

The importance of functional connectivity in the conservation of a ground-dwelling mammal in an urban Australian landscape

The distribution of the northern brown bandicoot (Isoodon macrourus), a medium-sized ground-dwelling marsupial, was examined in habitat fragments within the urban landscape of the city of Brisbane, Australia. From surveys conducted in 68 fragments, bandicoots were found to be present in 33 (49%) despite widespread habitat loss and fragmentation. Logistic regression analysis revealed that of 13 measured independent variables, functional connectivity was the only factor that significantly predicted the presence of bandicoots within fragments, with connectivity positively correlated with the likelihood of occupation. Functional connectivity was equated to the likelihood of bandicoot immigration into the focal fragment from the nearest occupied fragment, based on the estimated resistance to movement offered by the intervening matrix. Within Brisbane, riparian habitat fragments typically have a relatively high level of functional connectivity, as thin strips of vegetation fringing waterways serve as corridors between larger riparian areas and facilitate the movement of bandicoots between patches. Analyses based on the Akaike Information Criterion revealed that the optimal model based on landscape context variables was convincingly better supported by the data than the optimal model produced from fragment characteristics. However, it is important to examine both internal attributes of habitat fragments and external features of the surrounding landscape when modelling the distribution of ground-dwelling fauna in urban environments, or other landscapes with a highly variable matrix. As urban centres throughout the world expand, it is crucial that the ecology of local wildlife be considered to ensure functional connection is maintained between habitat patches, especially for the conservation of species that are highly susceptible to fragmentation.     

Forest cover and matrix functionality drive the abundance and reproductive success of an endangered primate in two fragmented rainforests

Landscape Ecology - Understanding population responses to landscape structure is critical to improve landscape planning. Yet, large uncertainty remains about how such responses vary among regions...     

Determinants of actual functional connectivity for calcareous grassland communities linked by rotational sheep grazing

In fragmented landscapes, plant species persistence depends on functional connectivity in terms of pollen flow to maintain genetic diversity within populations, and seed dispersal to re-colonize habitat patches following local extinction. Connectivity in plants is commonly modeled as a function of the physical distance between patches, without testing alternative dispersal vectors. In addition, pre- and post-dispersal processes such as seed production and establishment are likely to affect patch colonization rates. Here, we test alternative models of potential functional connectivity with different assumptions on source patch effects (patch area and species occupancy) and dispersal (relating to distance among patches, matrix composition, and sheep grazing routes) against empirical patch colonization rates at the community level (actual functional connectivity), accounting for post-dispersal effects in terms of structural elements providing regeneration niches for establishment. Our analyses are based on two surveys in 1989 and in 2009 of 48 habitat specialist plants in 62 previously abandoned calcareous grassland patches in the Southern Franconian Alb in Bavaria, Germany. The best connectivity model S _i , as identified by multi-model inference, combined distance along sheep grazing routes including consistently and intermittently grazed patches with mean species occupancy in 1989 as a proxy for pre-dispersal effects. Community-level patch colonization rates depended to equal degrees on connectivity and post-dispersal process. Our study highlights that actual functional connectivity of calcareous grassland communities cannot be approximated by structural connectivity based on physical distance alone, and modeling of functional connectivity needs to consider pre- and post-dispersal processes.     

Incorporating landscape elements into a connectivity measure: a case study for the Speckled wood butterfly (Pararge aegeria L.)

In spatial studies of populations, Euclidean distance is commonly used to measure the structural connectivity between habitat patches. The role of the matrix on patch connectivity is thereby ignored. However, the importance of the matrix for (dispersal) movement is increasingly being acknowledged. Our study compared the cost-distance measure with the Euclidean distance. The cost-distance is a simple GIS-calculated connectivity measure that incorporates the resistance of the landscape matrix to movement behaviour. We used presence-absence data from a field study on the Speckled wood butterfly in two Belgian landscapes. Logistic regression revealed that the cost-distance measure had a significantly better predictive power than the Euclidean distance. This result was consistent for all the six sets of different matrix resistance values. In our study the cost-distance proves to be a better connectivity measure than the Euclidean distance. This revised version was published online in July 2006 with corrections to the Cover Date.     

Landscape genetics matches with behavioral ecology and brings new insight on the functional connectivity in Mediterranean mouflon

Context

In natural populations, gene flow often represents a key factor in determining and maintaining genetic diversity. In a worldwide context of habitat fragmentation, assessing the relative contribution of landscape features to gene flow thus appears crucial for sustainable management of species.

Objective

We addressed this issue in Mediterranean mouflon (Ovis gmelini musimon?×?Ovis sp.) by combining previous knowledge on behavioral ecology with landscape genetics. We also assessed how sex-specific behavioral differences translated in term of functional connectivity in both sexes.

Methods

We relied on 239 individuals genotyped at 16 microsatellite markers. We applied a model optimization approach in a causal modeling framework of landscape genetics to test for the effects on gene flow of habitat types and linear landscape features previously identified as important for movements and habitat selection in both sexes. Five resistance values were alternately assigned to these landscape characteristics leading to a comprehensive set of resistance surfaces.

Results

Isolation by resistance shaped female gene flow, supporting the central role of linear landscape features as behavioral barriers for animal movements. Conversely, no isolation by resistance was detected in males. Although a lack of statistical power cannot be discarded to explain this result, it tended to confirm that males are less influenced by landscape structures during the mating period.

Conclusions

Combining previous knowledge on behavioral ecology with results from landscape genetics was decisive in assessing functional landscape connectivity in both sexes. These results highlighted the need to perform sex-specific studies for management and conservation of dimorphic species.

     

Quantifying functional connectivity: Experimental evidence for patch-specific resistance in the Natterjack toad (Bufo calamita)

Despite the importance assigned to inter-patch movements in fragmented systems, the structure of landscape between suitable habitat patches, the matrix, is often considered as to be of minor interest, or totally ignored. Consequently, models predicting metapopulation dynamics typically assume that dispersal and movement abilities are independent of the composition of the matrix. The predictions of such models should be invalided if that crucial assumption is unverified. In order to test the hypothesis of a patch-specific resistance, we led an experimental study to assess the matrix effects on the movement ability of juvenile Natterjack toads (Bufo calamita). The movement behaviour of first year toadlets, the dispersal stage in this species, was investigated in an arena experiment. Toadlet mobility was assessed in five landscape components that were mimicked in the lab: sandy soil, road, forest, agricultural field, and pasture. We analysed several movement components including move length, speed, efficiency and turning angle distribution. Our results showed that movement ability was strongly affected by the land cover, even if body size modulated the behavioural responses of toadlets. Performances were the best in the arenas mimicking sand and roads, and the worst in the forest arena, toadlet moves being three to five times less effective in the latter. The mobility was intermediate in the two other arenas. We propose here a new method to quantify functional connectivity, based on quantitative estimates of relative values for resistance of landscape components. This method offers a reliable alternative for resistance value estimates to subjective expert advice or inference from genetic population structure.     

Quantifying functional connectivity: Experimental evidence for patch-specific resistance in the Natterjack toad (Bufo calamita)

Despite the importance assigned to inter-patch movements in fragmented systems, the structure of landscape between suitable habitat patches, the matrix, is often considered as to be of minor interest, or totally ignored. Consequently, models predicting metapopulation dynamics typically assume that dispersal and movement abilities are independent of the composition of the matrix. The predictions of such models should be invalided if that crucial assumption is unverified. In order to test the hypothesis of a patch-specific resistance, we led an experimental study to assess the matrix effects on the movement ability of juvenile Natterjack toads (Bufo calamita). The movement behaviour of first year toadlets, the dispersal stage in this species, was investigated in an arena experiment. Toadlet mobility was assessed in five landscape components that were mimicked in the lab: sandy soil, road, forest, agricultural field, and pasture. We analysed several movement components including move length, speed, efficiency and turning angle distribution. Our results showed that movement ability was strongly affected by the land cover, even if body size modulated the behavioural responses of toadlets. Performances were the best in the arenas mimicking sand and roads, and the worst in the forest arena, toadlet moves being three to five times less effective in the latter. The mobility was intermediate in the two other arenas. We propose here a new method to quantify functional connectivity, based on quantitative estimates of relative values for resistance of landscape components. This method offers a reliable alternative for resistance value estimates to subjective expert advice or inference from genetic population structure.     

Improved connectivity analysis using multiple low-cost paths to evaluate habitat for the endangered San Martin titi monkey (Plecturocebus oenanthe) in north-central Peru

Context

Graph-theoretic evaluations of habitat connectivity often rely upon least-cost path analyses to evaluate connectedness of habitat patches, based on an underlying cost surface. We present two improvements upon these methods.

Objectives

As a case study to test these methods, we evaluated habitat connectivity for the endangered San Martin titi monkey (Plecturocebus oenanthe) in north-central Peru, to prioritize habitat patches for conservation.

Methods

First, rather than using a single least-cost path between habitat patches, we analyzed multigraphs made up of multiple low-cost paths. This allows us to differentiate between patches connected through a single narrow corridor, and patches connected by a wide swath of traversable land. We evaluate potential movement pathways by iteratively removing paths and recomputing connectivity metrics. Second, instead of performing a sensitivity analysis by varying costs uniformly across the landscape, we generated landscapes with spatially varying costs.

Results

This approach produced a more informative assessment of connectivity than standard graph analyses. Of the 4340 habitat patches considered across the landscape, we identified the most important 100, those frequently ranked highly through repeated network modifications, for multiple metrics and cost surfaces.

Conclusions

These methods represent a novel approach for assessing connectivity, better accounting for spatial configurations of habitat patches and uncertainty in cost surfaces. The ability to identify habitat patches with more possible routes to other patches is of interest for resiliency planning and prioritization in the face of continued habitat loss and climate change. These methods should be broadly applicable to conservation planning for other wildlife species.

     

Using animal movement behavior to categorize land cover and predict consequences for connectivity and patch residence times

Context

Landscape-scale population dynamics are driven in part by movement within and dispersal among habitat patches. Predicting these processes requires information about how movement behavior varies among land cover types.

Objectives

We investigated how butterfly movement in a heterogeneous landscape varies within and between habitat and matrix land cover types, and the implications of these differences for within-patch residence times and among-patch connectivity.

Methods

We empirically measured movement behavior in the Baltimore checkerspot butterfly (Euphydryas phaeton) in three land cover classes that broadly constitute habitat and two classes that constitute matrix. We also measured habitat preference at boundaries. We predicted patch residence times and interpatch dispersal using movement parameters estimated separately for each habitat and matrix land cover subclass (5 categories), or for combined habitat and combined matrix land cover classes (2 categories). We evaluated the effects of including edge behavior on all metrics.

Results

Overall, movement was slower within habitat land cover types, and faster in matrix cover types. Butterflies at forest edges were biased to remain in open areas, and connectivity and patch residence times were most affected by behavior at structural edges. Differences in movement between matrix subclasses had a greater effect on predictions about connectivity than differences between habitat subclasses. Differences in movement among habitat subclasses had a greater effect on residence times.

Conclusions

Our findings highlight the importance of careful classification of movement and land cover in heterogeneous landscapes, and reveal how subtle differences in behavioral responses to land cover can affect landscape-scale outcomes.

     

Linking movement behavior and fine-scale genetic structure to model landscape connectivity for bobcats (Lynx rufus)

Spatial heterogeneity can constrain the movement of individuals and consequently genes across a landscape, influencing demographic and genetic processes. In this study, we linked information on landscape composition, movement behavior, and genetic differentiation to gain a mechanistic understanding of how spatial heterogeneity may influence movement and gene flow of bobcats in the agricultural landscape of Iowa (USA). We analyzed movement paths of 23 animals to parameterize landscape resistance surfaces, applied least cost path analysis to generate measures of effective geographic distance between DNA collection locations of 625 bobcats, and tested the correlation between genetic distance and the different models of geographic distance. We found that bobcats showed a strong preference for forest over any other habitat type, and that incorporating information on habitat composition both along the path and in the surrounding landscape provided the best model of movement. Measures of effective geographic distance were significantly correlated with genetic distance, but not once the effects of Euclidean distance were accounted for. Thus, despite the impact of habitat composition on movement behavior, we did not detect a signature of a landscape effect in genetic structure. Our results are consistent with the issue of limiting factors: the high uniformity of forest fragmentation across southern Iowa, the primary study area, results in a landscape resistance pattern virtually indistinguishable from the isolation-by-distance pattern. The northern portion of the state, however, is predicted to pose a high level of resistance to bobcat movement, which may impede the regional genetic connectivity of populations across the Midwest.     

Long-term effects of managing for an endangered songbird on the heterogeneity of a fire-prone landscape

Context

Jack pine (Pinus banksiana)-dominated ecosystems of northern Lower Michigan are the primary breeding habitat for the federally endangered Kirtland’s warbler (Setophaga kirtlandii, KW). Historically, young stands used by KW were produced by stand-replacing wildfires, but fire suppression has necessitated the management of jack pine plantations for KW habitat since the 1970s. Effects of this long-term management on landscape age heterogeneity have previously not been quantified.

Objectives

We hypothesized that forest management has altered the spatial and temporal distribution of jack pine-dominated ecosystems beyond their historic range of variability.

Methods

By developing a diameter-age relationship for jack pine, we estimated ages of pre-European settlement trees found in General Land Office survey notes. We compared pre-European and current landscapes using geostatistical modeling of survey notes, and landscape metrics to quantify changes in pattern.

Results

Three KW management-based age classes (<20, 21–50, >50 years) are now more evenly distributed (31, 39, and 30 %, respectively) compared to the pre-European distribution (5, 19, 76 %) with little variability over time. Landscape metrics suggest the current landscape is younger and more fragmented than the pre-European landscape. These changes indicate restriction of the historic range of age variability, largely due to conversion of older jack pine stands to young KW habitat plantations.

Conclusions

Management has met KW population objectives, but has altered the temporal variability of the landscape’s age structure. Pre-European settlement patterns of stand-ages may provide a foundation for an ecosystem-based management plan for the region that supports both KW and the ecosystems upon which they depend.

     

Integrating animal movement with habitat suitability for estimating dynamic migratory connectivity

Context

High-resolution animal movement data are becoming increasingly available, yet having a multitude of empirical trajectories alone does not allow us to easily predict animal movement. To answer ecological and evolutionary questions at a population level, quantitative estimates of a species’ potential to link patches or populations are of importance.

Objectives

We introduce an approach that combines movement-informed simulated trajectories with an environment-informed estimate of the trajectories’ plausibility to derive connectivity. Using the example of bar-headed geese we estimated migratory connectivity at a landscape level throughout the annual cycle in their native range.

Methods

We used tracking data of bar-headed geese to develop a multi-state movement model and to estimate temporally explicit habitat suitability within the species’ range. We simulated migratory movements between range fragments, and calculated a measure we called route viability. The results are compared to expectations derived from published literature.

Results

Simulated migrations matched empirical trajectories in key characteristics such as stopover duration. The viability of the simulated trajectories was similar to that of the empirical trajectories. We found that, overall, the migratory connectivity was higher within the breeding than in wintering areas, corroborating previous findings for this species.

Conclusions

We show how empirical tracking data and environmental information can be fused for meaningful predictions of animal movements throughout the year and even outside the spatial range of the available data. Beyond predicting migratory connectivity, our framework will prove useful for modelling ecological processes facilitated by animal movement, such as seed dispersal or disease ecology.