• Barton K (2019) Package ‘MuMIn.’ R Packag version 1:
• Bartumeus F, Catalan J, Viswanathan GM, et al (2008) The influence of turning angles on the success of non-oriented animal searches. J Theor Biol 252:43–55. doi: 10.1016/j.jtbi.2008.01.009
• Bates D, Mächler M, Bolker B, Walker S (2015) Fitting Linear Mixed-Effects Models Using lme4. J Stat Softw 67:. doi: 10.18637/jss.v067.i01
• Bestley S, Jonsen ID, Hindell MA, et al (2012) Integrative modelling of animal movement: incorporating in situ habitat and behavioural information for a migratory marine predator. Proc R Soc B Biol Sci 280:20122262–20122262. doi: 10.1098/rspb.2012.2262
• Bonte D, Dahirel M (2017) Dispersal: a central and independent trait in life history. Oikos 126:472–479. doi: 10.1111/oik.03801
• Brakefield PM (1982a) Ecological Studies on the Butterfly Maniola jurtina in Britain. I. Adult Behaviour, Microdistribution and Dispersal. J Anim Ecol 51:713. doi: 10.2307/4000
• Brakefield PM (1982b) Ecological Studies on the Butterfly Maniola jurtina in Britain. II. Population Dynamics: The Present Position. J Anim Ecol 51:727. doi: 10.2307/4001
• Brown LM, Crone EE (2016a) Minimum area requirements for an at-risk butterfly based on movement and demography. Conserv Biol 30:103–112. doi: 10.1111/cobi.12588
• Brown LM, Crone EE (2016b) Individual variation changes dispersal distance and area requirements of a checkerspot butterfly. Ecology 97:106–115. doi: 10.1002/ecy.1216
• Chittka L, Dyer AG, Bock F, Dornhaus A (2003) Bees trade off foraging speed for accuracy. Nature 424:388–388. doi: 10.1038/424388a
• Conradt L, Bodsworth EJ, Roper TJ, Thomas CD (2000) Non-random dispersal in the butterfly Maniola jurtina: implications for metapopulation models. Proc R Soc B Biol Sci 267:1505–1510. doi: 10.1098/rspb.2000.1171
• Conradt L, Roper TJ (2006) Nonrandom movement behaviour at habitat boundaries in two butterfly species: Implications for dispersal. Ecology 87:125–132. doi: 10.1890/05-0413
• Conradt L, Roper TJ, Thomas CD (2001) Dispersal behaviour of individuals in metapopulations of two British butterflies. Oikos 95:416–424. doi: 10.1034/j.1600-0706.2001.950306.x
• Corvalan C, Hales S, McMichael AJ, et al (2005) Ecosystems and human well-being: health synthesis. World Health Organization
• Dapporto L, Habel JC, Dennis RLH, Schmitt T (2011) The biogeography of the western Mediterranean: elucidating contradictory distribution patterns of differentiation in Maniola jurtina (Lepidoptera: Nymphalidae). Biol J Linn Soc 103:571–577. doi: 10.1111/j.1095-8312.2011.01639.x
• Delattre T, Burel F, Humeau A, et al (2010) Dispersal mood revealed by shifts from routine to direct flights in the meadow brown butterfly Maniola jurtina. Oikos 119:1900–1908. doi: 10.1111/j.1600-0706.2010.18615.x
• Dennis RLH (1985) Polyommatus icarus on Brerteon Heath in Cheshire: voltinism and switches in resource exploitation. Ent Gaz 36:175–179
• Dennis RLH (1983) Egg-laying cues in the wall brown butterfly, Lasiommata megera (L.)(Lepidoptera: Satyridae). Entomol Gaz 34:89–95
• Dennis RLH (1992) Ecology of butterflies in Britain. Oxford University Press
• Dennis RLH (2012) A resource-based habitat view for conservation: butterflies in the British landscape. John Wiley & Sons
• Dover JW (1989) A method for recording and transcribing observations of butterfly behaviour. Entomol Gaz 40:95–100
• Dover JW, Clarke SA, Rew L (1992) Habitats and movement patterns of satyrid butterflies (Lepidoptera: Satyridae) on arable farmland. Entomol Gaz 43:29–44
• Dowdeswell W (1981) The life of the meadow brown, 1st edn. Butler & Tanner Ltd, London
• Dudley R, Srygley R (1994) Flight Physiology of Neotropical Butterflies: Allometry of Airspeeds During Natural Free Flight. J Exp Biol 191:125–39
• Dunning JB, Danielson BJ, Pulliam HR (1992) Ecological Processes That Affect Populations in Complex Landscapes. Oikos 65:169. doi: 10.2307/3544901
• Evans LC, Sibly RM, Thorbek P, et al (2019a) Integrating the influence of weather into mechanistic models of butterfly movement. Mov Ecol 7:24. doi: 10.1186/s40462-019-0171-7
• Evans LC, Sibly RM, Thorbek P, et al (2019b) Quantifying the effectiveness of agri-environment schemes for a grassland butterfly using individual-based models. Ecol Modell 411:. doi: 10.1016/j.ecolmodel.2019.108798
• Fahrig L, Merriam G (1994) Conservation of Fragmented Populations. Conserv Biol 8:50–59. doi: 10.1046/j.1523-1739.1994.08010050.x
• Fownes S, Roland J (2002) Effects of meadow suitability on female behaviour in the alpine butterfly Parnassius smintheus. Ecol Entomol 27:457–466. doi: 10.1046/j.1365-2311.2002.00426.x
• Fox R, Brereton TM, Asher J, et al (2015) The State of the UK’s Butterflies 2015
• Fred MS, O’Hara RB, Brommer JE (2006) Consequences of the spatial configuration of resources for the distribution and dynamics of the endangered Parnassius apollo butterfly. Biol Conserv 130:183–192. doi: 10.1016/j.biocon.2005.12.012
• Grant TJ, Parry HR, Zalucki MP, Bradbury SP (2018) 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. Ecol Modell 374:37–50
• Grimm V, Berger U, DeAngelis DL, et al (2010) The ODD protocol: A review and first update. Ecol Modell 221:2760–2768. doi: 10.1016/j.ecolmodel.2010.08.019
• Haddad NM (1999) Corridor Use Predicted from Behaviors at Habitat Boundaries. Am Nat 153:215–227. doi: 10.1086/303163
• Hanski I (1998) Metapopulation dynamics. Nature 396:41–49. doi: 10.1038/23876
• Haskell DG (1997) Experiments and a model examining learning in the area-restricted search behavior of ferrets ( Mustela putorius furo ). Behav Ecol 8:448–449. doi: 10.1093/beheco/8.4.448
• Heinz SK, Wissel C, Frank K (2006) The viability of metapopulations: Individual dispersal behaviour matters. Landsc Ecol 21:77–89. doi: 10.1007/s10980-005-0148-3
• Hovestadt T, Nowicki P (2008) Investigating Movement within Irregularly Shaped Patches: Analysis of Mark-Release-Recapture Data Using Randomization Procedures. Isr J Ecol Evol 54:137–154. doi: 10.1560/IJEE.54.1.137
• Ibrahim KM, Nichols RA, Hewitt GM (1996) Spatial patterns of genetic variation generated by different forms of dispersal during range expansion. Heredity (Edinb) 77:282–291. doi: 10.1038/hdy.1996.142
• Jackson HB, Fahrig L (2012) What size is a biologically relevant landscape? Landsc Ecol 27:929–941. doi: 10.1007/s10980-012-9757-9
• Jonsen ID, Myers RA, James MC (2006) Robust hierarchical state-space models reveal diel variation in travel rates of migrating leatherback turtles. J Anim Ecol 75:1046–1057. doi: 10.1111/j.1365-2656.2006.01129.x
• Kerr JT, Deguise I (2004) Habitat loss and the limits to endangered species recovery. Ecol Lett 7:1163–1169. doi: 10.1111/j.1461-0248.2004.00676.x
• Kokko H, López-Sepulcre A (2006) From individual dispersal to species ranges: Perspectives for a changing world. Science (80- ) 313:789–791. doi: 10.1126/science.1128566
• Kőrösi Á, Örvössy N, Batáry P, et al (2008) Restricted within-habitat movement and time-constrained egg laying of female Maculinea rebeli butterflies. Oecologia 156:455–464. doi: 10.1007/s00442-008-0986-1
• Kuznetsova A, Brockhoff PB, Christensen RHB (2017) lmerTest Package: Tests in Linear Mixed Effects Models. J Stat Softw 82:. doi: 10.18637/jss.v082.i13
• Lebeau J, Wesselingh RA, Van Dyck H (2017) Flower use of the butterfly Maniola jurtina in nectar-rich and nectar-poor grasslands: a nectar generalist with a strong preference? Insect Conserv Divers 10:258–270. doi: 10.1111/icad.12222
• Lebeau J, Wesselingh RA, Van Dyck H (2015) Butterfly density and behaviour in uncut hay meadow strips: Behavioural ecological consequences of an agri-environmental scheme. PLoS One 10:1–17. doi: 10.1371/journal.pone.0134945
• Lebeau J, Wesselingh RA, Van Dyck H (2016) Floral resource limitation severely reduces butterfly survival, condition and flight activity in simplified agricultural landscapes. Oecologia 180:421–427. doi: 10.1007/s00442-015-3492-2
• Lima SL, Zollner PA (1996) Towards a behavioral ecology of ecological landscapes. Trends Ecol Evol 11:131–5. doi: 10.1016/0169-5347(96)81094-9
• Lörtscher M (1997) Local movement patterns of three common grassland butterflies in a traditionally managed landscape. Bull Soc Entomol Suisse 70:43–55
• Lund U, Agostinelli C (2011) R package ‘circular’: Circular Statistics. URL https//r-forge r-project org/projects/circular
• Maier MJ (2014) DirichletReg: Dirichlet regression for compositional data in R
• Mair L, Thomas CD, Franco AMA, Hill JK (2015) Quantifying the activity levels and behavioural responses of butterfly species to habitat boundaries. Ecol Entomol 40:823–828. doi: 10.1111/een.12248
• Mason THE, Fortin D (2017) Functional responses in animal movement explain spatial heterogeneity in animal–habitat relationships. J Anim Ecol 86:960–971. doi: 10.1111/1365-2656.12682
• Morales JM, Ellner SP (2002) Scaling up animal movements in heterogeneous landscapes: the importance of behavior. Ecology 83:2240–2247
• Nakagawa S, Schielzeth H (2013) A general and simple method for obtaining R2 from generalized linear mixed‐effects models. Methods Ecol Evol 4:133–142
• Nathan R, Getz WM, Revilla E, et al (2008) A movement ecology paradigm for unifying organismal movement research. Proc Natl Acad Sci 105:19052–19059. doi: 10.1073/pnas.0800375105
• Nathan R, Klein EK, Robledo-Arnuncio JJ, Revilla E (2012) Dispersal kernels. In: Dispersal ecology and evolution. Oxford University Press Oxford, pp 187–210
• Norberg UM, Rayner JM V (1987) Ecological morphology and flight in bats (Mammalia; Chiroptera): wing adaptations, flight performance, foraging strategy and echolocation. Phil Trans R Soc Lond B 316:335–427
• Odendaal FJ, Turchin P, Stermitz FR, et al (1989) Influence of Host-Plant Density and Male Harassment on the Distribution of Female Euphydryas anicia (Nymphalidae ). 78:283–288
• Ouin A (2000) Hétérogénéité spatiale des paysages et processus écologiques: cas des rhopalocères dans le bocage breton
• Ouin A, Aviron S, Dover J, Burel F (2004) Complementation/supplementation of resources for butterflies in agricultural landscapes. Agric Ecosyst Environ 103:473–479. doi: 10.1016/j.agee.2003.11.003
• Ouin A, Martin M, Burel F (2008) Agricultural landscape connectivity for the meadow brown butterfly (Maniola jurtina). Agric Ecosyst Environ 124:193–199. doi: 10.1016/j.agee.2007.09.010
• Ovaskainen O, Hanski I (2004) From Individual Behavior to Metapopulation Dynamics: Unifying the Patchy Population and Classic Metapopulation Models. Am Nat 164:364–377. doi: 10.1086/423151
• Ovaskainen O, Luoto M, Ikonen I, et al (2008a) An Empirical Test of a Diffusion Model: Predicting Clouded Apollo Movements in a Novel Environment. Am Nat 171:610–619. doi: 10.1086/587070
• Ovaskainen O, Rekola H, Meyke E, Arjas E (2008b) Bayesian methods for analyzing movements in heterogeneous landscapes from mark-recapture data. Ecology 89:542–554. doi: 10.1890/07-0443.1
• Pauli BP, McCann NP, Zollner PA, et al (2013) SEARCH: Spatially Explicit Animal Response to Composition of Habitat. PLoS One 8:e64656. doi: 10.1371/journal.pone.0064656
• Pullin AS (1987) Changes in Leaf Quality Following Clipping and Regrowth of Urtica dioica, and Consequences for a Specialist Insect Herbivore, Aglais urticae. Oikos 49:39. doi: 10.2307/3565552
• Rákosy L, Schmitt T (2011) Are butterflies and moths suitable ecological indicator systems for restoration measures of semi-natural calcareous grassland habitats? Ecol Indic 11:1040–1045. doi: 10.1016/j.ecolind.2010.10.010
• Ravenscroft NOM (1994) The Ecology of the Chequered Skipper Butterfly Carterocephalus palaemon in Scotland. I. Microhabitat. J Appl Ecol 31:613. doi: 10.2307/2404152
• Reim E, Kahl S, Metschke K, Fischer K (2019) Sexual differences rather than flight performance underlie movement and exploration in a tropical butterfly. Ecol Entomol 44:648–658. doi: 10.1111/een.12738
• Roland J, Keyghobadi N, Fownes S (2000) Alpine Parnassius butterfly dispersal: Effects of landscape and population size. Ecology 81:1642–1653. doi: 10.1890/0012-9658(2000)081[1642:APBDEO]2.0.CO;2
• Root RB, Kareiva PM (1984) The search for resources by cabbage butterflies ( Pieris rapae): ecological consequences and adaptive significance of Markovian movements in a patchy environment. Ecology 65:147–165
• Saupe D (1988) Algorithms for random fractals. In: The science of fractal images. Springer, pp 71–136
• Scali V (1971) Imaginal Diapause and Gonadal Maturation of Maniola jurtina (Lepidoptera: Satyridae) from Tuscany. J Anim Ecol 40:467. doi: 10.2307/3255
• Schneider C (2003) The influence of spatial scale on quantifying insect dispersal: An analysis of butterfly data. Ecol Entomol 28:252–256. doi: 10.1046/j.1365-2311.2003.00495.x
• Schneider C, Dover J, Fry GLA (2003a) Movement of two grassland butterflies in the same habitat network: The role of adult resources and size of the study area. Ecol Entomol 28:219–227. doi: 10.1046/j.1365-2311.2003.00494.x
• Schneider C, Dover J, Fry GLA (2003b) Movement of two grassland butterflies in the same habitat network: the role of adult resources and size of the study area. Ecol Entomol 28:219–227
• Schtickzelle N, Joiris A, Dyck H Van, Baguette M (2007) Quantitative analysis of changes in movement behaviour within and outside habitat in a specialist butterfly. BMC Evol Biol 7:1–15. doi: 10.1186/1471-2148-7-4
• Schultz CB, Crone EE (2001) Edge-Mediated Dispersal Behavior in a Prairie Butterfly. Ecology 82:1879. doi: 10.2307/2680054
• Schultz CB, Dlugosch KM (1999) Nectar and hostplant scarcity limit populations of an endangered Oregon butterfly. Oecologia 119:231–238. doi: 10.1007/s004420050781
• Schultz CB, Franco AM a, Crone EE (2012) Response of butterflies to structural and resource boundaries. J Anim Ecol 81:724–734. doi: 10.1111/j.1365-2656.2011.01947.x
• Schultz CB, Haddad NM, Henry EH, Crone EE (2019a) Movement and Demography of At-Risk Butterflies: Building Blocks for Conservation. Annu Rev Entomol 64:167–184. doi: 10.1146/annurev-ento-011118-112204
• Schultz CB, Haddad NM, Henry EH, Crone EE (2019b) Movement and Demography of At-Risk Butterflies: Building Blocks for Conservation. Annu Rev Entomol 64:167–184. doi: 10.1146/annurev-ento-011118-112204
• Shreeve TG (1984) Habitat selection, mate location, and microclimatic constraints on the activity of the speckled wood butterfly Pararge aegeria. Oikos 42:371–377. doi: 10.2307/3544407
• Smith JNM (1974) The Food Searching Behaviour of Two European Thrushes. Behaviour 48:276–301. doi: 10.1163/156853974X00363
• Stevens VM, Turlure C, Baguette M (2010) A meta-analysis of dispersal in butterflies. doi: 10.1111/j.1469-185X.2009.00119.x
• Team RC (2018) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. 2017
• Thiele JC (2014) R Marries NetLogo : Introduction to the RNetLogo Package. J Stat Softw 58:1–41. doi: 10.18637/jss.v058.i02
• Thomas J. (2005) Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups. Philos Trans R Soc B Biol Sci 360:339–357. doi: 10.1098/rstb.2004.1585
• Thomas J (2010) Butterflies of britain and Ireland. British Wildlife Publishing
• Thomas JA (1983) The Ecology and Conservation of Lysandra bellargus (Lepidoptera: Lycaenidae) in Britain. J Appl Ecol 20:59. doi: 10.2307/2403376
• Thomas JA, Simcox DJ, Hovestadt T (2011) Evidence based conservation of butterflies. J Insect Conserv 15:241–258. doi: 10.1007/s10841-010-9341-z
• Thomas JA, Thomas CD, Simcox DJ, Clarke RT (1986) Ecology and Declining Status of the Silver-Spotted Skipper Butterfly (Hesperia comma) in Britain. J Appl Ecol 23:365. doi: 10.2307/2404023
• Tscharntke T, Tylianakis JM, Rand TA, et al (2012) Landscape moderation of biodiversity patterns and processes - eight hypotheses. Biol Rev 87:661–685. doi: 10.1111/j.1469-185X.2011.00216.x
• Turchin P (1991) Translating Foraging Movements in Heterogeneous Environments into the Spatial Distribution of Foragers. Ecology 72:1253–1266
• Turlure C, Van Dyck H (2009) On the consequences of aggressive male mate-locating behaviour and micro-climate for female host plant use in the butterfly Lycaena hippothoe. Behav Ecol Sociobiol 63:1581–1591. doi: 10.1007/s00265-009-0753-2
• Van Swaay C, Van Strien A, Harpke A, et al (2013) The European Grassland Butterfly Indicator: 1990–2011
• Van Swaay CAM, Dennis EB, Schmucki R, et al (2019) The EU Butterfly Indicator for Grassland species: 1990-2017: Technical Report. Butterfly Conserv Eur ABLE/eBMS
• Vickery JA, Feber RE, Fuller RJ (2009) Arable field margins managed for biodiversity conservation: A review of food resource provision for farmland birds. Agric Ecosyst Environ 133:1–13. doi: 10.1016/j.agee.2009.05.012
• Warren MS (1984) biology and status of the wood white butterfly, Leptidea sinapsis (L.)(Lepidoptera: Pieridae), in the British Isles. Entomol Gaz
• Wilensky U (1999) NetLogo. Evanston, IL: Center for connected learning and computer-based modeling, Northwestern University
• Wright DB, Herrington JA (2011) Problematic standard errors and confidence intervals for skewness and kurtosis. Behav Res Methods 43:8–17
• Zalucki MP, Kitching RL (1982) The Analysis and Description of Movement in Adult Danaus Plexippus L. (Lepidoptera: Danainae). Behaviour 80:174–197. doi: 10.1163/156853982X00346
• Zollner PA, Lima SL (2005) Behavioral tradeoffs when dispersing across a patchy landscape. Oikos 108:219–230