Functional composition and phenology of fruit-feeding butterflies in a fragmented landscape: variation of seasonality between habitat specialists

For butterflies, tolerance to the matrix may be an important criterion of habitat occurrence in fragmented landscapes. Here we examine the relative effects of habitat fragmentation and the surrounding agricultural matrix on the functional composition of fruit-feeding butterflies of the Atlantic rain forest in southeastern Brazil. Generalized linear models were used to detect the effects of landscape metrics on butterfly richness and abundance of the total assemblage and functional groups. Circular statistics were used to analyze the patterns of monthly abundance of the total assemblage and functional groups in the forest remnants and the surrounding matrices. In total, 650 butterflies representing 57 species were captured; species composition differed significantly between the forest fragments and the surrounding matrices. We recorded 22 forest specialists, 18 matrix specialists, 11 common species with matrix preference and six common species with forest preference. Forest connectivity favored the richness of forest specialists, while habitat fragmentation enhances the richness and abundance of matrix-tolerant species. Circular analysis revealed that forest specialists were more abundant in the rainy season while matrix-tolerant species proliferated in the dry season. Although maintaining connectivity of forest fragments may increase the mobility and dispersion of forest species, our results showed that landscape fragmentation modify butterfly assemblage by promoting an increase of matrix tolerant species with detriment of forest specialists.     

Landscape structure shapes habitat finding ability in a butterfly

Land-use intensification and habitat fragmentation is predicted to impact on the search strategies animals use to find habitat. We compared the habitat finding ability between populations of the speckled wood butterfly (Pararge aegeria L.) from landscapes that differ in degree of habitat fragmentation. Naïve butterflies reared under standardized laboratory conditions but originating from either fragmented agricultural landscapes or more continuous forested landscapes were released in the field, at fixed distances from a target habitat patch, and their flight paths were recorded. Butterflies originating from fragmented agricultural landscapes were better able to find a woodlot habitat from a distance compared to conspecifics from continuous forested landscapes. To manipulate the access to olfactory information, a subset of individuals from both landscape types were included in an antennae removal experiment. This confirmed the longer perceptual range for butterflies from agricultural landscapes and indicated the significance of both visual and olfactory information for orientation towards habitat. Our results are consistent with selection for increased perceptual range in fragmented landscapes to reduce dispersal costs. An increased perceptual range will alter the functional connectivity and thereby the chances for population persistence for the same level of structural connectivity in a fragmented landscape.     

A Flexible Approach for Assessing Functional Landscape Connectivity,with Application to Greater Sage-Grouse (Centrocercus urophasianus)

Connectivity of animal populations is an increasingly prominent concern in fragmented landscapes, yet existing methodological and conceptual approaches implicitly assume the presence of, or need for, discrete corridors. We tested this assumption by developing a flexible conceptual approach that does not assume, but allows for, the presence of discrete movement corridors. We quantified functional connectivity habitat for greater sage-grouse (Centrocercus urophasianus) across a large landscape in central western North America. We assigned sample locations to a movement state (encamped, traveling and relocating), and used Global Positioning System (GPS) location data and conditional logistic regression to estimate state-specific resource selection functions. Patterns of resource selection during different movement states reflected selection for sagebrush and general avoidance of rough topography and anthropogenic features. Distinct connectivity corridors were not common in the 5,625 km² study area. Rather, broad areas functioned as generally high or low quality connectivity habitat. A comprehensive map predicting the quality of connectivity habitat across the study area validated well based on a set of GPS locations from independent greater sage-grouse. The functional relationship between greater sage-grouse and the landscape did not always conform to the idea of a discrete corridor. A more flexible consideration of landscape connectivity may improve the efficacy of management actions by aligning those actions with the spatial patterns by which animals interact with the landscape.     

Functional connectivity as an indicator for patch occupancy in grassland specialists

Habitat specialists living in metapopulations are sensitive to habitat fragmentation. In most studies, the effects of fragmentation on such species are analyzed based on Euclidean inter-patch distances. This approach, however, ignores the role of the landscape matrix. Recently, therefore, functional distances that account for the composition of the landscape surrounding the habitat patches have been used more frequently as indicators for patch occupancy. However, the performance of functional and non-functional connectivity measures in predicting patch occupancy of such species has never been compared in a multi-species approach.Here we evaluate the effect of habitat connectivity on the patch occupancy of 13 habitat specialists from three different insect orders (Auchenorrhyncha, Lepidoptera, Orthoptera) in fragmented calcareous grasslands. In order to calculate functional distances we used four different sets of resistance values and rankings. We then modelled species’ occurrence using both Euclidean and functional (based on least-cost modelling) inter-patch distances as predictors.We found that functional connectivity measures provided better results than the non-functional approach. However, a functional connectivity measure that was based on very coarse land-cover data performed even better than connectivity measures that were based on much more detailed land-use data.In order to take into account possible effects of the landscape matrix on patch occupancy by habitat specialists, future metapopulation studies should use functional rather than Euclidean distances whenever possible. For practical applications, we recommend a ‘simple approach’ which requires only coarse land-cover data and in our study performed better than all other functional connectivity measures, even more complex ones.     

Dispersal in the Glanville fritillary butterfly in fragmented versus continuous landscapes: comparison between three methods

1. Habitat fragmentation may lead to natural selection on dispersal rate and other life‐history traits. Both theoretical analyses and empirical studies suggest that habitat fragmentation may select either for increased or decreased dispersal depending on the traits of the species and the characteristics of the landscape. 2. Dispersal and movement rates in Glanville fritillary butterflies (Melitaea cinxia) originating from a continuous landscape in China and from a highly fragmented landscape in Finland were compared using three different methods. 3. The methods included replicated mark‐release‐recapture (MRR) experiments conducted in the natural environments in China and Finland, tracking with harmonic radar of captive‐reared but free‐flying butterflies in a common environment in the field, and replicated common garden experiments in a large outdoor population cage. 4. The results were largely consistent, showing that butterflies from the more continuous landscape in China had a lower movement rate than butterflies originating from the fragmented landscape in Finland. Butterflies originating from newly‐established populations in Finland moved significantly longer distances than butterflies originating from old populations in Finland or from China, demonstrating significant intra‐specific variation in dispersal rate in Finland. These results are consistent with model predictions for the Glanville fritillary. 5. The tracking experiment revealed a result that would have been impossible to obtain with MRR experiments: movement rate was influenced by a significant interaction between population origin (China vs. Finland) and ambient air temperature.     

Gene flow and functional connectivity in the natterjack toad

Functional connectivity is a key factor for the persistence of many specialist species in fragmented landscapes. However, connectivity estimates have rarely been validated by the observation of dispersal movements. In this study, we estimated functional connectivity of a real landscape by modelling dispersal for the endangered natterjack toad (Bufo calamita) using cost distance. Cost distance allows the evaluation of 'effective distances', which are distances corrected for the costs involved in moving between habitat patches in spatially explicit landscapes. We parameterized cost-distance models using the results of our previous experimental investigation of natterjack's movement behaviour. These model predictions (connectivity estimates from the GIS study) were then confronted to genetic-based dispersal rates between natterjack populations in the same landscape using Mantel tests. Dispersal rates between the populations were inferred from variation at six microsatellite loci. Based on these results, we conclude that matrix structure has a strong effect on dispersal rates. Moreover, we found that cost distances generated by habitat preferences explained dispersal rates better than did the Euclidian distances, or the connectivity estimate based on patch-specific resistances (patch viscosity). This study is a clear example of how landscape genetics can validate operational functional connectivity estimates.     

Barriers to dispersal of rain forest butterflies in tropical agricultural landscapes

Fragmentation of natural habitats can be detrimental for species if individuals fail to cross habitat boundaries to reach new locations, thereby reducing functional connectivity. Connectivity is crucial for species shifting their ranges under climate change, making it important to understand factors that might prevent movement through human‐modified landscapes. In tropical regions, rain forests are being fragmented by agricultural expansion, potentially isolating populations of highly diverse forest‐dependent species. The likelihood of crossing habitat boundaries is an important determinant of species dispersal through fragmented landscapes, and so we examined movement across rain forest‐oil palm plantation boundaries on Borneo by using relatively mobile nymphalid butterflies as our model study taxon. We marked 1666 individuals from 65 species, and 19 percent (100/527) of recaptured individuals crossed the boundary. Boundary crossing was relatively frequent in some species, and net movement of individuals was from forest into plantation. However, boundary crossing from forest into plantation was detected in less than 50 percent (12/28) of recaptured species and was dominated by small‐sized butterfly species whose larval host plants occurred within plantations. Thus, while oil palm plantations may be relatively permeable to some species, they may act as barriers to the movement of forest‐dependent species (i.e., species that require rain forest habitat to breed), highlighting the importance of maintaining forest connectivity for conserving rain forest species.     

The effect of an agro-pasture landscape on diversity and migration patterns of frugivorous butterflies in Chiapas, Mexico

Few relevant data are available to analyze how landscape structure and composition affect the abundance and movement patterns of tropical insects. Using mark-release recapture experiments we examine the effect of an agro-pasture matrix on changes in diversity and migration of frugivorous butterflies in a tropical fragmented landscape in southern Mexico. In total, 53 frugivorous butterfly species were recorded in the entire landscape. Butterfly species composition was much more similar between sites than plant composition. A total of 3,501 individuals belonging to 41 butterfly species were captured, out of which 23 species (56%) were recaptured at least once. A large fraction of individuals was recaptured at the site of release (91%). We failed to find a significant relationship between the proportion occupied by the matrix and rates of residence, emigration, and immigration. Our results suggest that matrix quality in this and other traditionally managed agro-pasture landscapes plays a key role in both keeping important levels of biodiversity and maintaining constant movements of butterflies between otherwise isolated habitat patches.     

Effects of Wildflower Strip Quality,Quantity, and Connectivity on Butterfly Diversity in a Swiss Arable Landscape

The effects of habitat restoration measures designated to promote farmland biodiversity have been documented at the field scale, but little is known about their role in restoring the agricultural mosaic. In this study, we analyzed the effects of wildflower strips (WFS) at the field scale and in the landscape context on butterflies in a Swiss arable landscape. Three hypotheses were tested: (1) butterfly diversity and abundance are higher in WFS than in conventional fields; (2) butterfly diversity and abundance are enhanced by the amount, proximity and connectivity of WFS in the landscape context; (3) additional factors influence butterfly diversity and abundance according to individual site conditions and landscape context characteristics linked to other landscape elements. WFS had more species and individuals of butterflies than conventional habitats. However, promoted species were mainly generalists; few specialists were enhanced. The diversity of all butterflies and of generalists increased linearly with percent cover of WFS, reflecting an effect of restoration measures depending on the landscape context. The influence of proximity and connectivity of WFS were, however, not significant. The occurrence of specialists was conditioned by plant species richness, while the effect of WFS for overall diversity was affected by the amount of grassland in the surroundings. We conclude that to increase the effectiveness of biodiversity‐orientated restoration measures, their implementation should be steered toward increasing the share of WFS in the landscape. However, the combination of WFS with additional restoration measures might be needed to halt the decline of specialist species.     

Assessing the landscape functional connectivity using movement maps: a case study with endemic Azorean insects

There is a vast body of literature aiming to predict, for a large number of taxa, the spatial distribution of suitable areas given the expected future changes of climatic conditions. However, such studies often overlook the role of landscape functional connectivity. This is particularly relevant for species with low vagility, as ground-dwelling insects, inhabiting areas with high human pressure due to habitat destruction and fragmentation, namely in the islands. In this study, we developed an individual-based model (IBM) that simulates individual movement according to landscape resistance and mortality probability, in order to derive the landscape movement map, and applied it to five endemic ground-dwelling insects of Terceira Island (Azores). We then confronted the movement maps of each species against the species distribution models previously developed for both current and future climatic conditions, quantifying the amount of important movement areas that are enclosed by the distribution polygons. We further sought to identify where habitat restoration would increase the overall connectivity among large habitat patches. Our results showed that, for both timeframes, the distribution models enclosed small amounts of areas predicted to be important for animal movement. Additionally, we predicted strong reductions (up to 94%) of these important areas for functional connectivity. We also identified areas in-between native forest of primary importance for restoration that may significantly increase the probability of persistence of our model species. We anticipate that this study will be useful to both conservation planners and ecologists seeking to understand species movement and dispersal both is islands and elsewhere.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

The Influence of Landscape Grain Size on Butterfly Diversity in Grasslands

The relationship between butterfly diversity and both habitat and landscape variables was studied in two areas of southern Sweden. The habitat quality of the grasslands was similar in the two study areas but the landscape pattern differed in grain size and amount of grassland and forest. Using a transect survey method, a total of 3341 butterflies were observed and 30 taxa identified. We found that both habitat and landscape variables influenced the butterfly diversity of the investigated grasslands. Species composition differed markedly between the two study areas. A study area with a fine-grained landscape pattern, a high cover of semi-natural grassland and many forest edges had twice as many butterfly species but half the number of individuals compared with a coarser-grained study area with larger grasslands widely spread in a matrix of arable fields. The results of our study indicate that both habitat quality and landscape pattern have to be considered when developing conservation strategies for grassland butterflies.     

Fruit‐feeding Butterfly Communities are Influenced by Restoration Age in Tropical Forests

Currently, a large‐scale restoration project aims to restore around 15 million hectares of Atlantic Forest in Brazil. This will increase forest cover and connectivity among remnant sites as well as restore environmental services. Currently, studies on recovery of fauna in restored areas of the Atlantic Forest are practically nonexistent. To address this knowledge vacuum, our study compares diversity patterns of fruit‐feeding butterflies in three forest areas with different restoration ages (11, 22, and 54 years), and uses a native forest area as reference. Results showed butterfly communities in maturing restored areas becoming more similar to the ones found in the native forest, with an increase in the proportional abundance of forest species, and a decrease of edge and grassland species. Moreover, we found a higher diversity among sites at the intermediate restoration age, with a community composed of both grassland and forest species. Butterfly species composition differed significantly among sites, showing interesting patterns of potential species replacement over time. Our results indicate that, although restored sites were located in a fragmented landscape, they provide suitable habitats for recolonization by fruit‐feeding butterfly assemblages. Hence, restored areas can be considered important habitat for forest animal species, increasing local biodiversity and, possibly, restoring some of the ecosystem services provided by them.     

Transcriptome Analysis Reveals Signature of Adaptation to Landscape Fragmentation

We characterize allelic and gene expression variation between populations of the Glanville fritillary butterfly (Melitaea cinxia) from two fragmented and two continuous landscapes in northern Europe. The populations exhibit significant differences in their life history traits, e.g. butterflies from fragmented landscapes have higher flight metabolic rate and dispersal rate in the field, and higher larval growth rate, than butterflies from continuous landscapes. In fragmented landscapes, local populations are small and have a high risk of local extinction, and hence the long-term persistence at the landscape level is based on frequent re-colonization of vacant habitat patches, which is predicted to select for increased dispersal rate. Using RNA-seq data and a common garden experiment, we found that a large number of genes (1,841) were differentially expressed between the landscape types. Hexamerin genes, the expression of which has previously been shown to have high heritability and which correlate strongly with larval development time in the Glanville fritillary, had higher expression in fragmented than continuous landscapes. Genes that were more highly expressed in butterflies from newly-established than old local populations within a fragmented landscape were also more highly expressed, at the landscape level, in fragmented than continuous landscapes. This result suggests that recurrent extinctions and re-colonizations in fragmented landscapes select a for specific expression profile. Genes that were significantly up-regulated following an experimental flight treatment had higher basal expression in fragmented landscapes, indicating that these butterflies are genetically primed for frequent flight. Active flight causes oxidative stress, but butterflies from fragmented landscapes were more tolerant of hypoxia. We conclude that differences in gene expression between the landscape types reflect genomic adaptations to landscape fragmentation.     

Mapping functional connectivity

An objective and reliable assessment of wildlife movement is important in theoretical and applied ecology. The identification and mapping of landscape elements that may enhance functional connectivity is usually a subjective process based on visual interpretations of species movement patterns. New methods based on mathematical morphology provide a generic, flexible, and automated approach for the definition of indicators based on the classification and mapping of spatial patterns of connectivity from observed or simulated movement and dispersal events. The approach is illustrated with data derived from simulated movement on a map produced from satellite imagery of a structurally complex, multi-habitat landscape. The analysis reveals critical areas that facilitate the movement of dispersers among habitat patches. Mathematical morphology can be applied to any movement map providing new insights into pattern-process linkages in multi-habitat landscapes.     

The taste of nectar – a neglected area of pollination ecology

We investigated the impact of landscape structure on landscape connectivity using a combination of simulation and empirical experiments. In a previous study we documented the movement behaviour of a specialized goldenrod beetle ( Trirhabda borealis Blake) in three kinds of patches: habitat (goldenrod) patches and two types of matrix patch (cut vegetation and cut vegetation containing camouflage netting as an impediment to movement). In the current study, we used this information to construct simulation and experimental landscapes consisting of mosaics of these three patch types, to study the effect of landscape structure on landscape connectivity, using the T. borealis beetle as a model system. In the simulation studies, landscape connectivity was based on movements of individual beetles, and was measured in six different ways. The simulations revealed that the six measures of landscape connectivity were influenced by different aspects of landscape structure, suggesting that: (1) landscape connectivity is a poorly defined concept, and (2) the same landscape may have different landscape connectivity values when different measures of landscape connectivity are used. There were two general predictions that held over all measures of landscape connectivity: (1) increasing interpatch distance significantly decreased landscape connectivity and (2) the influence of matrix elements on landscape connectivity was small in comparison to the influence of habitat elements. Empirical mark-release-resight experiments using Trirhabda beetles in experimental landscapes supported the simulation results.     

An empirical test of a diffusion model: predicting clouded apollo movements in a novel environment

Functional connectivity is a fundamental concept in conservation biology because it sets the level of migration and gene flow among local populations. However, functional connectivity is difficult to measure, largely because it is hard to acquire and analyze movement data from heterogeneous landscapes. Here we apply a Bayesian state-space framework to parameterize a diffusion-based movement model using capture-recapture data on the endangered clouded apollo butterfly. We test whether the model is able to disentangle the inherent movement behavior of the species from landscape structure and sampling artifacts, which is a necessity if the model is to be used to examine how movements depend on landscape structure. We show that this is the case by demonstrating that the model, parameterized with data from a reference landscape, correctly predicts movements in a structurally different landscape. In particular, the model helps to explain why a movement corridor that was constructed as a management measure failed to increase movement among local populations. We illustrate how the parameterized model can be used to derive biologically relevant measures of functional connectivity, thus linking movement data with models of spatial population dynamics.     

Where movement happens: scale‐dependent landscape effects on genetic differentiation in the European tree frog

Functional connectivity among fragmented populations depends on the landscape matrix between occupied habitat patches and its effect on the frequency of animal movement and gene flow. The quantification of landscape effects on gene flow should therefore be scale‐dependent. Here, we explored the impact of different spatial scales in a landscape genetic analysis of the European tree frog Hyla arborea in a fragmented landscape in Switzerland. We examined the effects of landscape elements and geographic distance on genetic differentiation at three distance classes reflecting varying frequencies of tree frog movement. We calculated pairwise F_ST‐values and assembled 16 landscape elements within 1 km wide corridors between all pairs of tree frog breeding sites. Per distance class, we computed a multiple regression model with stepwise backward elimination and permutation testing. At distances of<2 km, only a larger river acted as a barrier to gene flow. At distances>2 km, geographic distance had a negative effect on gene flow as had landscape elements such as forests and roads. In general, hedgerows and various structure‐rich landscape elements positively affected gene flow. As we found distinct scale‐dependent landscape effects on gene flow, future landscape genetic studies should analyse the effects of landscape variables at different spatial dimensions relevant for the movement and dispersal of the study organisms. Corresponding studies should also carefully consider relevant correlations among the landscape elements tested and should preferentially replicate their analysis at the landscape‐level in order to avoid idiosyncratic results owing to the particular scale and landscape studied.     

Preliminary Information on the Home Range and Movement Patterns of Sturnira lilium (Phyllostomidae) in a Naturally Fragmented Landscape in Bolivia

Here, we used radiotelemetry to determine home range and movement patterns of the frugivorous bat Sturnira lilium in a naturally fragmented landscape in Beni, Bolivia. Four females were monitored for up to 7 d. Based on 575 radiotelemetry positions, kernel home ranges were between 36.5 and 90.7 ha. During each night, bats visited most of their home range, which included forest islands and continuous forest; however activity was concentrated in core-use areas representing 6–12 percent of the home ranges. There was moderate overlap in home range and core-use areas among the four females. The mean long axis across the kernel home ranges was 1324 m. The results reported here show that S. lilium is able to move freely among forest elements in the landscape while crossing a savanna matrix. Hence, S. lilium may contribute to maintain connectivity and ecological processes in a fragmented landscape.     

High and dry or sunk and dunked: lessons for tallgrass prairies from quaking bogs

Northern Wisconsin bogs provide a natural experiment on butterfly population occurrence in a naturally highly fragmented vegetation type, which may provide insight on conserving butterflies in anthropogenically fragmented and degraded landscapes. We surveyed butterflies in bogs (about as unaffected by humans as possible, but naturally occurring over <1% of northern Wisconsin) primarily during 2002?C2009, with additional observations from 1986 to 2001. Different bog types had different bog-specialist butterfly faunas, but bog butterfly abundance also differed in similar vegetations among subregions. Some small isolated bogs held very high densities of specialist butterflies. Summer but not spring specialists frequented adjacent lowland roadsides and utilized a variety of non-native as well as native nectar sources. Paleo-entomology indicates that insects don??t evolve out of trouble; instead they move out of trouble. Given the low dispersal apparent today for species restricted to bogs, ??move?? might be better understood as ??hunkering?? within their vegetation as it expands and shrinks and moves around the landscape. Although bogs appeared to have more intact specialist butterfly faunas than tallgrass prairies (99.9% destroyed by human activities), bog butterflies do not live in average sites even in a relatively natural landscape. Just as bog butterflies are ??sunk and dunked?? in isolation, specialist butterflies elsewhere may have been left ??high and dry?? naturally, or are now due to human activities. Numerous studies have demonstrated that presence and abundance of specialist butterflies increase with increasing size and connectedness of habitat patches. But with long-term consistent vegetation, populations with high abundances in small isolated sites and with low numbers thinly occurring in large sites can be secure, as shown by bog butterflies.