Butterfly Movement Research Articles

Caterpillar movement mediates spatially local interactions and determines the relationship between population density and contact.

While interactions in nature are inherently local, ecological models often assume homogeneity across space, allowing for generalization across systems and greater mathematical tractability. Density-dependent disease models are a prominent example of models that assume homogeneous interactions, leading to the prediction that disease transmission will scale linearly with population density. In this study, we examined how the scale of larval butterfly movement interacts with the resource landscape to influence the relationship between larval contact and population density in the Baltimore checkerspot (Euphydryas phaeton). Our study was inspired by the recent discovery of a viral pathogen that is transmitted horizontally among Baltimore checkerspot larvae. We used multi-year larvae location data across six Baltimore checkerspot populations in the eastern U.S. to test whether larval nests are spatially clustered. We then integrated these spatial data with larval movement data in different resource contexts to investigate whether heterogeneity in spatially local interactions alters the assumed linear relationship between larval nest density and contact. We used Correlated Random Walk (CRW) models and field observations of larval movement behavior to construct Probability Distribution Functions (PDFs) of larval dispersal, and calculated the overlap in these PDFs to estimate conspecific contact within each population. We found that all populations exhibited significant spatial clustering in their habitat use. Subsequent larval movement rates were influenced by encounters with host plants and larval age, and under many movement scenarios, the scale of predicted larval movement was not sufficient to allow for the "homogeneous mixing" assumed in density dependent disease models. Therefore, relationships between population density and larval contact were typically non-linear. We also found that observed use of available habitat patches led to significantly greater contact than would occur if habitat use were spatially random. These findings strongly suggest that incorporating larval movement and spatial variation in larval interactions is critical to modeling disease outcomes in E. phaeton. Epidemiological models that assume a linear relationship between population density and larval contact have the potential to underestimate transmission rates, especially in small populations that are already vulnerable to extinction.

Biological Aspects and Movements of Neotropical Fruit-Feeding Butterflies.

The patterns of insect movement are the cumulate product of many individual decisions and can be shaped by the way morphology and behaviour interacts with landscape structure and composition. Hence, the ongoing process of habitat fragmentation increases the distance among suitable habitats and changes those in such a way that it may favour some movement behaviour. Our study described some biological traits (sex ratio, age structure and individual permanence in a population), as well as the movements of fruit-feeding butterflies along the horizontal dimension (among habitats: forest interior, natural forest transition - ecotone and anthropogenic forest transition - edge) and the vertical dimension (between canopy and understory). We sampled butterflies monthly over 1year in the Atlantic rainforest, South-eastern Brazil, following a standardized design using bait traps. We found that sex ratio was male biased for five out of the six more abundant species and the age structure showed an increase in recruitment of new individuals in the dry season followed by a noticeable aging of the populations in the wet season. In general, our results revealed an aggregated spatial distribution, in which few individuals travelled long distances while most individuals were recaptured in the same trap, suggesting that all studied habitats currently provide the necessary conditions for the maintenance of butterfly populations, favouring fewer movements and narrow home ranges for both sexes and species. Conservation of a set of heterogeneous habitats it is especially important for the maintenance of sedentary butterflies and of those that move large distances.

Comparison between butterfly optimization algorithm and particle swarm optimization for tuning cascade PID control system of PMDC motor

In this paper, two optimization methods are used to adjust the gain values for the cascade PID controller. These algorithms are the butterfly optimization algorithm (BOA), which is a modern method based on tracking the movement of butterflies to the scent of a fragrance to reach the best position and the second method is particle swarm optimization (PSO). The PID controllers in this system are used to control the position, velocity, and current of a permanent magnet DC motor (PMDC) with an accurate tracking trajectory to reach the desired position. The simulation results using the Matlab environment showed that the butterfly optimization algorithm is better than the particle swarming optimization (PSO) in terms of performance and overshoot or any deviation in tracking the path to reach the desired position. While an overshoot of 2.557% was observed when using the PSO algorithm, and a position deviation of 7.82 degrees was observed from the reference position.

Path-planning in 3D space using butterfly optimization algorithm

The Butterfly Optimization Algorithm is one of the most recent nature-inspired algorithms that mimic the butterflies' behavior in mating and finding food, for solving the global optimization problems. The algorithm utilizes the sense of butterflies of smelling for determining the location of nectar and find mates, which is based on the foraging strategy of those insects. This paper represents a method of using the BOA algorithm for solving the problem of path planning in three-dimensional space. The proposed method finds a path from a particular starting point to any chosen goal, where the generated final path is completely safe and collision-free. The algorithm is based on 3 phases: the initial phase, the iteration phase, and the final phase. The movement of butterflies is based on two search moves, one of them is Local random search; where the butterfly moves randomly within the swarm, and the other is Global search; where the butterfly moves towards the best-fitted butterfly in the current population. The proposed method in this paper is able to find a collision-free path from the start point to the goal in all of the presented test environments in proximately well performance and the results were computed in terms of execution time and path length.

Behavioural modes in butterflies: their implications for movement and searching behaviour

Animals move in ‘modes’ where movement patterns relate to specific behaviours. Despite much work on the movement of butterflies, their behavioural modes are relatively unexplored. Here we analysed the behaviour of the model butterfly species the meadow brown, Maniola jurtina. We identified modes in both sexes and across habitats varying in resource density. We found that, in nectar-rich habitats, males had more diverse behaviour than females, engaging in a unique ‘high-flight’ mode associated with mate search, whereas females were primarily nectaring or inactive. In nectar-poor habitats, both sexes were similar, switching between flight and inactivity. We also identified the movement parameters of the modes, finding that, for both sexes, movements associated with nectaring were slower and more tortuous and, for males, the mode associated with mate searching was straighter and faster. Using an individual-based random-walk model, we investigated the effects of behaviour on movement predictions by comparing a mode-switching model with a version including intraspecific variation and another assuming homogeneity between individuals. For both sexes, including modes affected the mean and shape of the displacement rate compared to models assuming homogeneity, although for females modes increased displacement 1.5 times while for males they decreased it by a third. Both models also differed substantially from models assuming intraspecific variation. Finally, using a new model of search behaviour we investigated the general conditions under which individuals should engage in an exclusive search for host plants or receptive females. Parameterized for M. jurtina, the model predicted males should engage exclusively in mate search, but females only when searching is very efficient. The model provides a framework for analysing the searching behaviour of other butterfly species.

The importance of including habitat-specific behaviour in models of butterfly movement

Dispersal is a key process affecting population persistence and major factors affecting dispersal rates are the amounts, connectedness and properties of habitats in landscapes. We present new data on the butterfly Maniola jurtina in flower-rich and flower-poor habitats that demonstrates how movement and behaviour differ between sexes and habitat types, and how this effects consequent dispersal rates. Females had higher flight speeds than males, but their total time in flight was four times less. The effect of habitat type was strong for both sexes, flight speeds were ~ 2.5 × and ~ 1.7 × faster on resource-poor habitats for males and females, respectively, and flights were approximately 50% longer. With few exceptions females oviposited in the mown grass habitat, likely because growing grass offers better food for emerging caterpillars, but they foraged in the resource-rich habitat. It seems that females faced a trade-off between ovipositing without foraging in the mown grass or foraging without ovipositing where flowers were abundant. We show that taking account of habitat-dependent differences in activity, here categorised as flight or non-flight, is crucial to obtaining good fits of an individual-based model to observed movement. An important implication of this finding is that incorporating habitat-specific activity budgets is likely necessary for predicting longer-term dispersal in heterogeneous habitats, as habitat-specific behaviour substantially influences the mean (> 30% difference) and kurtosis (1.4 × difference) of dispersal kernels. The presented IBMs provide a simple method to explicitly incorporate known activity and movement rates when predicting dispersal in changing and heterogeneous landscapes.

Behavior underpins the predictive power of a trait-based model of butterfly movement.

Dispersal ability is key to species persistence in times of environmental change. Assessing a species' vulnerability and response to anthropogenic changes is often performed using one of two methods: correlative approaches that infer dispersal potential based on traits, such as wingspan or an index of mobility derived from expert opinion, or a mechanistic modeling approach that extrapolates displacement rates from empirical data on short‐term movements.Here, we compare and evaluate the success of the correlative and mechanistic approaches using a mechanistic random‐walk model of butterfly movement that incorporates relationships between wingspan and sex‐specific movement behaviors.The model was parameterized with new data collected on four species of butterfly in the south of England, and we observe how wingspan relates to flight speeds, turning angles, flight durations, and displacement rates.We show that flight speeds and turning angles correlate with wingspan but that to achieve good prediction of displacement even over 10 min the model must also include details of sex‐ and species‐specific movement behaviors.We discuss what factors are likely to differentially motivate the sexes and how these could be included in mechanistic models of dispersal to improve their use in ecological forecasting.

Permeability of habitat edges for Ringlet butterflies (Lepidoptera, Nymphalidae, Erebia Dalman 1816) in an alpine landscape

We tracked the movements of adult Ringlet butterflies (Lepidoptera, Nymphalidae, Erebia Dalman, 1816) in high-elevation (> 1800 meters a.s.l.) grasslands in the Austrian Alps in order to test if an anthropogenic boundary (= an asphalt road) had a stronger effect on butterfly movement than natural habitat boundaries (trees, scree, or dwarf shrubs surrounding grassland sites). 373 individuals (136 females, 237 males) belonging to 11 Erebia species were observed in one flight season (July–August 2013) while approaching or crossing habitat edges. Erebia pandrose (Borkhausen, 1788) was the most abundant species with 239 observations. All species studied were reluctant to cross habitat boundaries, but permeability was further strongly affected by the border type. Additional variables influencing movement probability were species identity and the time of the day. In E. pandrose, for which we had sufficient observations to analyse this, individuals were more likely to cross a boundary in the morning and in the late afternoon than at midday. Erebia euryale (Esper, 1805) and E. nivalis Lorković & de Lesse, 1954 were more likely to leave a habitat patch than their studied congeners. The key result of our study is that the paved road had the lowest permeability among all edge types (0.1 likelihood of crossing when approaching the edge). A road cutting across a conservation area (viz. a national park) thus hinders inter-patch exchange among Ringlet butterflies in the alpine zone, even though theoretically they ought to be able to fly across.

Integrating the influence of weather into mechanistic models of butterfly movement

BackgroundUnderstanding the factors influencing movement is essential to forecasting species persistence in a changing environment. Movement is often studied using mechanistic models, extrapolating short-term observations of individuals to longer-term predictions, but the role of weather variables such as air temperature and solar radiation, key determinants of ectotherm activity, are generally neglected. We aim to show how the effects of weather can be incorporated into individual-based models of butterfly movement thus allowing analysis of their effects.MethodsWe constructed a mechanistic movement model and calibrated it with high precision movement data on a widely studied species of butterfly, the meadow brown (Maniola jurtina), collected over a 21-week period at four sites in southern England. Day time temperatures during the study ranged from 14.5 to 31.5 °C and solar radiation from heavy cloud to bright sunshine. The effects of weather are integrated into the individual-based model through weather-dependent scaling of parametric distributions representing key behaviours: the durations of flight and periods of inactivity.ResultsFlight speed was unaffected by weather, time between successive flights increased as solar radiation decreased, and flight duration showed a unimodal response to air temperature that peaked between approximately 23 °C and 26 °C. After validation, the model demonstrated that weather alone can produce a more than two-fold difference in predicted weekly displacement.ConclusionsIndividual Based models provide a useful framework for integrating the effect of weather into movement models. By including weather effects we are able to explain a two-fold difference in movement rate of M. jurtina consistent with inter-annual variation in dispersal measured in population studies. Climate change for the studied populations is expected to decrease activity and dispersal rates since these butterflies already operate close to their thermal optimum.

Movement and Demography of At-Risk Butterflies: Building Blocks for Conservation.

The number of insect species at risk of population decline and extinction is increasing rapidly. Yet we know almost nothing about the ecology of these species, except for at-risk butterflies. A growing body of literature shows how butterfly vital rates, including demography and movement, are essential for guiding conservation and recovery. History has shown us that without these data, conservation decisions often weaken, rather than enhance, population viability. This is especially true in changing landscapes. We review knowledge of vital rates across all at-risk butterflies. We have information on movement for 17 of 283 butterfly species and information on demography for 19 species. We find that habitat-specific movement behavior is key to understanding how to connect populations, and habitat-specific demography is central to managing habitats. Methods and analyses worked out for butterflies can provide a scaffold around which to build studies for the conservation of other at-risk insects.

Predicting monarch butterfly (Danaus plexippus) movement and egg-laying with a spatially-explicit agent-based model: The role of monarch perceptual range and spatial memory

An ∼80% decline in the eastern population of the monarch butterfly (Danaus plexippus) has prompted conservation efforts to increase summer reproductive success in the Midwest United States. Implementation of conservation practices will create a patchwork of milkweed (mainly Asclepias spp.) habitat within agricultural landscapes dominated by corn and soybean production. Since the monarch butterfly is a vagile species, reproductive success is, in part, a function of both the amount and spatial arrangement of habitat patches in a fragmented landscape. To inform conservation planning we developed a spatially-explicit, agent-based model for summer breeding, non-migratory female monarch butterfly movement and egg-laying on an Iowa, USA landscape. Our model employs a unique movement algorithm when monarch agents encounter habitat edges that incorporates monarch perceptual range to their host plant and spatial memory of previously visited habitat. These behavioral factors are rarely incorporated into animal movement algorithms; however, they can influence estimates of resource utilization. Model exploration assessed the distribution and density of eggs laid on a spatially-explicit 148,665 ha landscape comprised of 17 land cover classes with varying milkweed densities. Uncertainty analysis was undertaken by sampling 25 combinations of perceptual range, spatial memory, flight step length and flight directionality parameters from a total of 256 (44) possible combinations. Movement paths simulated with our new movement algorithm show preferential use of high density milkweed areas that would not be simulated using a correlated random walk. Increasing perceptual range caused a decrease in the area used by monarch agents and caused a skewed egg distribution where most eggs were laid in relatively few habitat patches. Increasing spatial memory caused an increase in the area used but decreased the median number of eggs laid in roadside habitat. Current national and regional monarch conservation goals assume a uniform distribution of milkweed in different land cover classes. Translating these goals into spatially-explicit, heterogeneous habitat patches is essential for predicting realized fecundity in the landscape. Our model provides the foundation to link national and regional monarch conservation goals to fine scale spatial configurations of habitat patches in defined landscapes.

Impacts of solar energy on butterfly communities in mediterranean agro‐ecosystems

The rapid expansion of Utility‐Scale Solar Energy (USSE) is expected to meet economic and environmental challenges that society faces today and in the future. Yet there is a paucity of comprehensive research on biodiversity responses to USSE. Here we investigated the impact of USSE on species movements via the correlations between landscape connectivity modeling and the similarity of butterfly communities given their life‐history traits. Our results suggest that mobile butterflies cope with USSE while interpatch movement of sedentary butterflies does not depend on landscape structure. We provide land‐managers and energy‐developers with an innovative tool to assess the ecological integration of USSE. © 2017 American Institute of Chemical Engineers Environ Prog, 36: 1817–1823, 2017

Fine Scale Movements of the Butterfly Plebejus argus in a Heterogeneous Natural Landscape as Revealed by GPS Tracking

The study of butterfly movements has focused on dispersal behaviour in the framework of population persistence in heterogeneous landscapes. The ecological significance of routine movements has received less attention. These movements may be influenced by structural attributes of habitat patches or may reflect the distribution of food, mates, host plants or ecological interactions. The relative influence of structural and functional factors on flight patterns is poorly understood, partly because butterfly movements are often described by simplified representations of actual trajectories. Using high-resolution GPS tracking we obtained accurate trajectories of routine movements of Plebejus argus in a heterogeneous natural landscape. Habitat quality in patches was ranked according to the abundance of host and nectar plants as well as the abundance of nests of its mutualistic ant Lasius niger. Movements were slow and winding in high quality habitats whereas faster, straighter flights were observed in poor habitats. At edges, butterflies often crossed without any exploratory behaviour towards patches of better quality, suggesting they may use cues to detect resources at some distance. Conversely, individuals usually stayed in the patch after exploring edges with other patches of lower quality. However, scanning also preceded exits towards clearly unsuitable habitat, compatible with transfers to distant high-quality patches. We conclude that patterns of movement in P. argus were explained by spatial heterogeneity defined by functional rather than structural criteria. We also show that inexpensive handheld GPS receivers allow depicting detailed flying trajectories in open flat terrain revealing complex behavioural patterns.

A case study of butterfly road kills from Anaikatty Hills, Western Ghats, Tamil Nadu, India

<p><strong> </strong>Anaikatty Hills of the Western Ghats in Tamil Nadu witness the annual spectacle of mass movement of lakhs of butterflies. The present paper examines the impact of vehicular traffic on this ‘butterfly migration’ through a survey of butterfly mortality along a road stretch in Anaikatty Hills. A high rate of mortality due to road traffic was observed during the mass movement of butterflies. One-hundred-and-thirty-five butterfly road kills belonging to three families, nine genera and 12 species were recorded during the study. The proportion of nymphalid butterflies among the road kills (70%) was very high compared to their respective share in the background population (39%), indicating a higher road mortality risk for nymphalids. The conservation significance of the road traffic impact on butterfly assemblage and management options are discussed.</p><div> </div>

Fine-Grained Distribution of a Non-Native Resource Can Alter the Population Dynamics of a Native Consumer.

New interactions with non-native species can alter selection pressures on native species. Here, we examined the effect of the spatial distribution of a non-native species, a factor that determines ecological and evolutionary outcomes but that is poorly understood, particularly on a fine scale. Specifically, we explored a native butterfly population and a non-native plant on which the butterfly oviposits despite the plant’s toxicity to larvae. We developed an individual-based model to describe movement and oviposition behaviors of each butterfly, which were determined by plant distribution and the butterfly's host preference genotype. We estimated the parameter values of the model from rich field data. We simulated various patterns of plant distributions and compared the rates of butterfly population growth and changes in the allele frequency of oviposition preference. Neither the number nor mean area of patches of non-native species affected the butterfly population, whereas plant abundance, patch shape, and distance to the nearest native and non-native patches altered both the population dynamics and genetics. Furthermore, we found a dramatic decrease in population growth rates when we reduced the distance to the nearest native patch from 147 m to 136 m. Thus changes in the non-native resource distribution that are critical to the fate of the native herbivore could only be detected at a fine-grained scale that matched the scale of a female butterfly’s movement. In addition, we found that the native butterfly population was unlikely to be rescued by the exclusion of the allele for acceptance of the non-native plant as a host. This study thus highlights the importance of including both ecological and evolutionary dynamics in analyses of the outcome of species interactions and provides insights into habitat management for non-native species.

Low relative error in consumer-grade GPS units make them ideal for measuring small-scale animal movement patterns.

Consumer-grade GPS units are a staple of modern field ecology, but the relatively large error radii reported by manufacturers (up to 10 m) ostensibly precludes their utility in measuring fine-scale movement of small animals such as insects. Here we demonstrate that for data collected at fine spatio-temporal scales, these devices can produce exceptionally accurate data on step-length and movement patterns of small animals. With an understanding of the properties of GPS error and how it arises, it is possible, using a simple field protocol, to use consumer grade GPS units to collect step-length data for the movement of small animals that introduces a median error as small as 11 cm. These small error rates were measured in controlled observations of real butterfly movement. Similar conclusions were reached using a ground-truth test track prepared with a field tape and compass and subsequently measured 20 times using the same methodology as the butterfly tracking. Median error in the ground-truth track was slightly higher than the field data, mostly between 20 and 30 cm, but even for the smallest ground-truth step (70 cm), this is still a signal-to-noise ratio of 3:1, and for steps of 3 m or more, the ratio is greater than 10:1. Such small errors relative to the movements being measured make these inexpensive units useful for measuring insect and other small animal movements on small to intermediate scales with budgets orders of magnitude lower than survey-grade units used in past studies. As an additional advantage, these units are simpler to operate, and insect or other small animal trackways can be collected more quickly than either survey-grade units or more traditional ruler/gird approaches.

Development and Verification of a Protocol to Quantify Hip Joint Kinematics

Background: The butterfly save technique is commonly used by ice hockey goaltenders and has recently been identified as a potential mechanism for hip joint injuries due to the extreme body positions involved. Unfortunately, commonly used kinematic marker sets that determine these body positions are heavily influenced by skin motion artifact and are obscured by protective equipment, making it difficult to obtain reliable measures of hip motion. Purpose: To create a new kinematic protocol that could be used when protective equipment prevents typical marker placements and to use this protocol to quantify hip kinematics and butterfly performance in 4 different goalie pad conditions. Study Design: Controlled laboratory study. Methods: A new marker set consisting of marker clusters attached to the lateral thigh and posterior leg was developed. This marker set was verified by evaluating the root mean square (RMS) difference between the developed testing marker set and the calibrated anatomic systems technique marker set during passive range of motion (ROM) tests. The testing marker set was then used in a repeated-measures study in which 12 junior goaltenders performed 5 butterfly movements on synthetic ice, in 4 different goalie pad conditions (control, flexible-wide leg channel, flexible-tight leg channel, and stiff-wide leg channel). Results: The grouped RMS differences and SDs calculated during verification were 1.43° ± 0.41°, 1.0° ± 0.39°, and 3.32° ± 1.32° for hip flexion-extension, abduction-adduction, and internal-external rotation, respectively. There was no significant main effect of goal pad condition on the peak amount of hip internal rotation; however, there was a significant main effect of goal pad condition on the butterfly width (P = .022). Post hoc comparisons revealed that the butterfly width was significantly smaller in the control pad condition compared with the flexible-tight pad condition (P = .03). Conclusion: The new marker set enabled measurements of hip joint kinematics while subjects are wearing protective equipment that are not possible with other marker sets. Interindividual variations in performance of the butterfly technique influenced the amount of hip internal rotation achieved; however, on average, goaltenders exceeded their active internal ROM during butterfly movements. Clinical Relevance: Exceeding internal rotation range of hip motion may make goaltenders susceptible to hip injuries such as femoral acetabular impingement.

Demography and mobility of three common understory butterfly species from tropical rain forest of Papua New Guinea

AbstractThe mobility of butterflies determines their ability to find host plant species, and thus their potential host plant range, as well as their ability to maintain meta‐populations in fragmented habitats. While butterfly movement has been extensively studied for temperate species, very little is known for tropical forest species. A mark‐release‐recapture study of the three most common butterfly species in the understory of a lowland primary rainforest in Papua New Guinea included 3,705, 394 and 317 marked individuals of Danis danis, Taenaris sp. and Parthenos aspila respectively, with 1,031, 78 and 40 butterfly individuals recaptured at least once. Over a period of 6 weeks there were almost 22,000 individuals belonging to these three species hatching within or entering our four study plots totaling 14.58 ha in area. The most abundant species, D. danis, with 20,000 individuals, showed highly variable population densities during the study. The residency time in the studied plots was highest for P. aspila (84 days), as individual butterflies stayed mostly in a single gap; we estimated that less than 1 % of individuals disperse 1 km or more. Similar movement probability was found in D. danis whilst in Taenaris sp., 10 % of the population disperses ≥1 km. Movement distances of D. danis were more than sufficient to locate its host plant, Derris elliptica, which occurred in 61 % of the 20 × 20 m subplots within a 50 ha plot. Compared with temperate species, our three species have much longer life spans, but their movement patterns remain within the known mobility estimates of temperate species. The mobility of D. danis is close to the average for temperate Lycaenidae, while Taenaris sp. is more mobile and P. aspila less mobile than the mean for all temperate species.

Kinematic Analysis of Butterfly Swimming

This study aimed to examine the relative phase of the arm and leg movements and the percentage of each of the 4 arm movement phases at three race paces of the butterfly swimming between two levels of swimmers based on the temporal analysis. Eight elite male swimmers and eight club level college male swimmers participated in the study. Two underwater high-speed cameras (200Hz) were set in the transverse and sagittal plane under the water and synchronized to capture one cycle of butterfly stroke movement. The kinematics data were digitized and calculated with the Kwon 3D software. The continuous relative phase was calculated using the normalized time series of the angular displacement and angular velocity of the shoulder and knee joints angles. Macroscopically, one cycle of the arm-leg coordination pattern can be observed between two levels at three race paces in butterfly stroke: a consistent relationship (2:1) exists between two dolphin kicks and one arm stroke in butterfly swimming. The continuous relative phase analyses showed that, in the beginning of the stroke-catch phase, the continuous relative phase was from anti-phase (-180°) to in-phase (50°). For the pull phase, the in-phase relation (±50°) remained unchanged throughout the stage. During the push phase, the relative phase gradually increased toward anti-phase (180°). The percentage of catch, pull, push, and recovery phases were not evenly distributed across one cycle of the butterfly stroke. Especially in elite group, there is a trend of decreasing the percentage of catch phase. In conclusion, these results provide useful information for swimmers as well as coaches to examine the swimming movement of the butterfly stroke.

Changes in butterfly movements along a gradient of land use in farmlands of Transylvania (Romania)

Context Agricultural transformation and increased land use intensity often lead to simplified landscapes and biodiversity loss. For animals, one possible mechanism underpinning biodiversity loss in agricultural landscapes is the disruption of movements. The disruption of movements may explain, for example, why butterfly communities in agricultural landscapes are often dominated by generalist species with high mobility. Objectives Here, we investigated how the movement patterns of butterflies characterised by different levels of mobility changed along a gradient of agricultural land use intensity. Methods To this end, we studied 15 landscapes in low-intensity farmland in Central Romania, measuring 10 ha each and covering a gradient of landscape heterogeneity and woody vegetation cover. In these landscapes, we tracked movements of 563 individuals of nine butterfly species. Results Our findings showed that overall movement activities differed significantly between species, corresponding well with expert-derived estimates of species-specific mobility. Interestingly, species of low and high mobility responded in opposite ways to increasing levels of landscape heterogeneity. In relatively simple landscapes, the movement patterns of low and high mobility species were similar. By contrast, in complex landscapes, the flight paths of low-mobility species became shorter and more erratic, whereas the flight paths of high-mobility species became longer and straighter. An analysis of the land covers traversed showed that most species avoided arable land but favoured the more heterogeneous parts of a given landscape. Conclusions In combination, our results suggest that non-arable patches in agricultural landscapes are important for butterfly movements, especially for low-mobility species.