With ongoing insect declines, species expanding in distribution and abundance deserve attention, as understanding their success may help design conservation strategies for less successful species. Common causes of these successes include warmer climates, novel resources, and exploiting land use change, including land abandonment. These factors affect the nymphalid butterfly Neptis rivularis, developing on Spiraea spp. shrubs and reaching the north-western limits of its trans-Palearctic distribution in Central Europe. We combined mark-recapture, behaviour analysis, and distribution modelling to study N. rivularis in wetlands of the Třeboňsko Protected Landscape (IUCN category V). The long-living adults (up to 4 weeks) spent a considerable amount of time searching for partners, ovipositing and nectaring at Spiraea shrubs, alternating this with stays in tree crowns, where they located cool shelters, spent nights, and presumably fed on honeydew. They formed high-density populations (310 adults/ha), exploiting high host plant abundance. They adhered to floodplains and to conditions of relatively mild winters. The ongoing Spiraea encroachment of abandoned alluvial grasslands is, thus, a transient situation, ultimately followed by forest encroachment. Rewilding the habitats by introducing native ungulates presents an opportunity to restore the disturbance regime of the sites. The increased resource supply combined with a warming climate has opened up temperate Europe to colonization by N. rivularis.

Faculty of Agrobiology Food and Natural Resources Kamýcká 129 165 00 Praha Suchodl Czech Republic

Faculty of Nature and Technology Hochschule Bremen City University of Applied Sciences Neustadtswall 30 28199 Bremen Germany

Faculty of Science University of South Bohemia Branisovska 31 37005 Ceske Budejovice Czech Republic

Institute of Entomology Biological Centre CAS Branisovska 31 37005 Ceske Budejovice Czech Republic

Muzeum Regionu Valašsko 755 01 Vsetín Czech Republic

See more in PubMed

Albrecht J, Bureš J, Cepák J, Čapek M, Havlíček J, Hora J, Klimeš Z, Kloubec B, Kubelka V., Pakandl M, Pecl K, Pelz P, Pykal J, Šťastný K (2015) Ptáci jižních Čech [Birds of South Bohemia]. Jihočeský kraj, České Budějovice.

Auckland JN, Debinski DM, Clark WR (2004) Survival, movement, and resource use of the butterfly Parnassius clodius. Ecol Entomol 29:139–149. https://doi.org/10.1111/j.0307-6946.2004.00581.x DOI

Bárta D, Dolný A (2007) Vážky České republiky: Ekologie, ochrana a rozšíření [Odonata of the Czech Republic: ecology, conservation and distribution]. ČSOP Vlašim, Vlašim

Battaglia LL, Minchin PR, Pritchett DW (2002) Sixteen years of old-field succession and reestablishment of a bottomland hardwood forest in the Lower Mississippi Alluvial Valley. Wetlands 22:1–17. https://doi.org/10.1672/0277-5212(2002)022[0001:SYOOFS]2.0.CO;2 DOI

Beck J, Mühlenberg E, Fiedler K (1999) Mud-puddling behavior in tropical butterflies: in search of proteins or minerals? Oecologia 119:140–148. https://doi.org/10.1007/s004420050770 PubMed DOI

Benes J, Konvicka M, Dvorak J, Fric Z, Havelda Z, Pavlicko A, Vrabec V, Weidenhoffer Z (2002) MotýliČeské republiky rozšíření a ochrana I, II. [Butterflies of the Czech Republic: distribution and conservation I, II]. SOM, Prague

Bhatade R, Patwardhan A, Kurve PN (2019) A preliminary study on some ecological factors involved in mud-puddling of butterflies in Mumbai metropolitan region. Int J Entomol Res 4:1–6. https://doi.org/10.13140/RG.2.2.13854.46401 DOI

Błoński W, Gwardjan M (2016) Występowanie pasyna lucylla Neptis rivularis (Scopoli, 1763) (Lepidoptera: Nymphalidae) i jego rośliny żywicielskiej – tawuły wierzbolistnej Spiraea salicifolia Linnaeus, 1753 (Rosaceae) w okolicach Starachowic [Occurrence of the Hungarian glider Neptis rivularis (Scopoli, 1763) (Lepidoptera: Nymphalidae) and its food plant – bridewort Spiraea salicifolia Linnaeus, 1953 (Rosaceae) near Starachowice]. Naturalia 5:126–131

Bohnet IC, Molnarova KJ, van den Brink A, Beilin R, Sklenicka P (2022) How cultural heritage can support sustainable landscape development: the case of Třeboň Basin, Czech Republic. Landsc Urban Plan 226:104492. https://doi.org/10.1016/j.landurbplan.2022.104492 DOI

Bonari G, Fajmon K, Malenovský I, Zelený D, Holuša J, Jongepierová I, Kočárek P, Konvička O, Uřičář J, Chytrý M (2017) Management of semi-natural grasslands benefiting both plant and insect diversity: the importance of heterogeneity and tradition. Agric Ecosyst Environ 246:243–252. https://doi.org/10.1016/j.agee.2017.06.010 DOI

ter Braak CJF, Šmilauer P (2018) Canoco reference manual and user’s guide: software for ordination (version 5.10). Biometris, Wageningen University & Research

Burkart B, Konold W, Gärtner M (2003) Der Einfluss von Schafen, Ziegen und Elchen auf die Vegetation des ehemaligen Panzerschießplatzes Dauban. In: Konold W, Burkart B (eds) Offenland Und Naturschutz. Schriftenreihe Des Instituts Für Landespflege Der AlbertLudwigs-Universität, Freiburg, Culterra, pp 217–234

Bury J (2020) Common Glider-Neptis sappho (Pallas, 1771) (Lepidoptera: Nymphalidae) in south-eastern Poland: another case of oversight or rapid range expansion of a species considered extinct. Ann Upper Silesian Mus Bytom Entomol 29:1–20. https://doi.org/10.5281/zenodo.4066591 DOI

Cannon RJC (2019) Courtship and mating in butterflies. CABI Books, Wallingford

Carlson CJ (2020) embarcadero: species distribution modelling with Bayesian additive regression trees in r. Methods Ecol Evol 11:850–858. https://doi.org/10.1111/2041-210X.13389 DOI

Carver S, Convery I, Hawkins S, Beyers R, Eagle A, Kun Z, van Maanen E, Cao Y, Fisher M, Edwards SR, Nelson C, Gann GD, Shurter S, Aguilar K, Andrade A, Ripple WJ, Davis J, Sinclair A, Bekoff M, Noss R, Foreman D, Pettersson H, Root-Bernstein M, Svenning JC, Taylor P, Wynne-Jones S, Featherstone AW, Flojgaard C, Stanley-Price M, Navarro LM, Aykroyd T, Parfitt A, Soule M (2021) Guiding principles for rewilding. Conserv Biol 35:1882–1893. https://doi.org/10.1111/cobi.13730 PubMed DOI

Chytrý M, Tichý L (2018) National vegetation classification of the Czech Republic: a summary of the approach. Phytocenologia 48:121–131. https://doi.org/10.1127/phyto/2017/0184 DOI

Clench HK (1966) Behavioral thermoregulation in butterflies. Ecology 47:1021–1034. https://doi.org/10.2307/1935649 DOI

Cooch EG, White GC (2022) Program MARK: a gentle introduction (21 ed.). http://www.phidot.org/software/mark/docs/book/

Cook LM, Brower LP, Croze HJ (1967) The accuracy of a population estimation from multiple recapture data. J Anim Ecol 36:57–60. https://doi.org/10.2307/3014 DOI

Corke D (1999) Are honeydew/sap-feeding butterflies (Lepidoptera: Rhopalocera) affected by particulate air-pollution? J Insect Conserv 3:5–14. https://doi.org/10.1023/A:1009670404398 DOI

Crisan A, Sitar C, Craioveanu MC, Vizauer TC, Rakosy L (2014) Multiannual population size estimates and mobility of the endemic Pseudophilotes bavius hungarica (Lepidoptera: Lycaenidae) from Transylvania (Romania). North-Western J Zool 10:S115–S124

Davies ZG, Wilson RJ, Coles S, Thomas CD (2006) Changing habitat associations of a thermally constrained species, the silver-spotted skipper butterfly, in response to climate warming. J Anim Ecol 75:247–256. https://doi.org/10.1111/j.1365-2656.2006.01044.x PubMed DOI

Dennis RLH (ed) (1992) The ecology of butterflies in Britain. Oxford University Press, Oxford

Dolek M, Geyer A (1997) Influence of management on butterflies of rare grassland ecosystems in Germany. J Insect Conserv 1:125–130. https://doi.org/10.1023/A:1018443412153 DOI

Duncan P (1982) Horses and grasses. The nutritional ecology of equids and their impact on the Camargue. Ecological Studies 87, Springer, New York

Dziekańska I, Nowicki P, Pirożnikow E, Sielezniew M (2020) A unique population in a unique area: the Alcon Blue butterfly and its specific parasitoid in the Białowieża Forest. Insects 11:687. https://doi.org/10.3390/insects11100687 PubMed DOI PMC

Ehl S, Holzhauer SIJ, Ryrholm N, Schmitt T (2019) Phenology, mobility and behaviour of the arcto-alpine species Boloria napaea in its arctic habitat. Sci Rep 9:1–9. https://doi.org/10.1038/s41598-019-40508-7 DOI

Ehrlich PR (1965) The population biology of the butterfly, Euphydryas editha. II. The structure of the Jasper Ridge colony. Evolution 19:327–336. https://doi.org/10.2307/2406442 DOI

Eliot JN (1969) An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Bull Brit Mus (Natural History) Entomol Suppl. 15:1–155

Fagerstrom T, Wiklund C (1982) Why do males emerge before females – protandry as a mating strategy in male and female butterflies. Oecologia 52(2):164–166 PubMed DOI

Fayet CMJ, Reilly KH, Van Ham C, Verburg PH (2022) What is the future of abandoned agricultural lands? A systematic review of alternative trajectories in Europe. Land Use Policy 112:105833. https://doi.org/10.1016/j.landusepol.2021.105833 DOI

Ferster B, Vulinec K (2010) Population size and conservation of the last eastern remnants of the regal fritillary, Speyeria idalia (Drury) [Lepidoptera, Nymphalidae]; implications for temperate grassland restoration. J Insect Conserv 14:31–42. https://doi.org/10.1007/s10841-009-9222-5 DOI

Fiedler K (1993) The remarkable biology of two Malaysian lycaenid butterflies. Nature Malaysiana 18:35–43

Fischer MK, Volkl W, Hoffmann KH (2005) Honeydew production and honeydew sugar composition of polyphagous black bean aphid, Aphis fabae (Hemiptera: Aphididae) on various host plants and implications for ant-attendance. Eur J Entomol 102:155–160. https://doi.org/10.14411/eje.2005.025 DOI

Fric Z, Konvicka M (2007) Dispersal kernels of butterflies: power-law functions are invariant to marking frequency. Basic Appl Ecol 8:377–386. https://doi.org/10.1016/j.baae.2006.06.005 DOI

Garrido P, Naumov V, Söderquist L, Jansson A, Thulin C-G (2022) Effects of experimental rewilding on butterflies, bumblebees and grasshoppers. J Insect Conserv 26:763–771. https://doi.org/10.1007/s10841-022-00420-4 DOI

Geier T (2016) Beobachtungen zum Auftreten des Arealerweiterers Pieris mannii (Mayer, 1851) im Gebiet der unteren Nahe (Rheinland-Pfalz) mit Nachweisen dreier Raupennahrungspflanzen im Freiland (Lepidoptera: Pieridae). Nachr Entomol Ver Apollo 37:27–40

Gorbunov PY (2001) The butterflies of Russia: classification, genitalia, keys for identification. Thesis, Ekaterinburg.

Haase VH, Brunk I, Voigtländer K, Balkenhol B (2019) Invasion of the neophyte Spiraea sp. and succession of species-rich grasslands: impacts on the communities of spiders (Araneae), ground beetles (Carabidae), millipedes (Diplopoda) and centipedes (Chilopoda). Ber Naturfors Ges Oberlausitz 27:5–42

Habel JC, Schmitt T, Gros P, Ulrich W (2022) Breakpoints in butterfly decline in Central Europe over the last century. Sci Tot Environ 851:158315. https://doi.org/10.1016/j.scitotenv.2022.158315 DOI

Habel JC, Eberle J, Charo J, Maghenda M, Schmitt T (2023) Population ecology and behaviour of two Afrotropical forest butterflies. J Insect Conserv 27:1–11. https://doi.org/10.1007/s10841-022-00451-x DOI

Hanč Z, Beneš J, Faltynek Fric Z, Pavlíčko A, Zapletal M (2019) Denní motýli a vřetenušky jižních Čech [Butterflies and burnet moths of South Bohemia]. Jihočeský kraj, České Budějovice

Hardy PB, Dennis RLH (2004) Seasonal and daily bias in butterfly perch height and substrate choice. Entomol Gaz 55:239–248. https://doi.org/10.1007/s10905-007-9073-4 DOI

Hejda R, Farkač J, Chobot K (eds) (2017) Červený seznam ohrožených druhů České republiky. Bezobratlí. [Red List of Threatened Species of the Czech Republic. Invertebrates]. Příroda, 36:1–612

Hijmans RJ, Williams E, Vennes C, Hijmans MRJ (2017) Package ‘geosphere’. Spherical trigonometry 1:1–45

Hill JK, Thomas CD, Lewis OT (1996) Effects of habitat patch size and isolation on dispersal by Hesperia comma butterflies: implications for metapopulation structure. J Anim Ecol 65:725–735. https://doi.org/10.2307/5671 DOI

Hlásek J, Hlásek O (1997) Ze života bělopáska tavolníkového [On the life of the Hungarian glider]. Živa 45:173–174

Hönigová J, Chobot K (2014) Jemné předivo české krajiny v GIS: konsolidovaná vrstva ekosystémů [Fine fabric of Czech landscape in GIS: consolidated ecosystems layer]. Ochrana Přírody 69:27–30

Höttinger H, Pendl M, Wiemers M, Pospisil A (2013) Insekten in Wien - Tagfalter. Österreichische Gesellschaft für Entomofaunistik, Vienna, p 347

Janík T, Peters W, Šálek M, Romportl D, Jirků M, Engleder T, Ernst M, Neudert J, Heurich M (2021) The declining occurrence of moose (Alces alces) at the southernmost edge of its range raise conservation concerns. Ecol Evol 11:5468–5483. https://doi.org/10.1002/ece3.7441 PubMed DOI PMC

Joyce CB (2014) Ecological consequences and restoration potential of abandoned wet grasslands. Ecol Eng 66:91–102. https://doi.org/10.1016/j.ecoleng.2013.05.008 DOI

Jugovic J, Uboni C, Zupan S, Lužnik M (2018) Demography of the endangered butterfly Euphydryas aurinia (Lepidoptera: Nymphalidae): a case study of populations in sub-Mediterranean dry calcareous grasslands. Eur J Entomol 115:493–503. https://doi.org/10.14411/eje.2018.049 DOI

Jutzeler D (1983) Beobachtungen über Neptis rivularis Scop. im Tessin. Mitt Entomol Ges Basel 33:10–14

Jutzeler D, Hoettinger H, Malicky M, Rebeusek F, Sala G, Verovnik R (2000) Biology of Neptis sappho (Pallas, 1771) based on the monograph by Timpe & Timpe (1993) and its actual distribution and conservation status in Austria, Italy and Slovenia (Lepidoptera: Nymphalidae). Linneana Belgica 17:315–330

Karger D, Conrad O, Böhner J, Kawohl T, Kreft H, Soria-Auza RW, Zimmermann NE, Linder HP, Kessler M (2017) Climatologies at high resolution for the Earth’s land surface areas. Sci Data 4:170122. https://doi.org/10.1038/sdata.2017.122 PubMed DOI PMC

Keenleyside C, Tucker GM (2010) Farmland abandonment in the EU: an assessment of trends and prospects. Report for WWF, Institute for European Environmental Policy, London

Kemp DJ (2001) Investigating the consistency of mate-locating behavior in the territorial butterfly Hypolimnas bolina (Lepidoptera: Nymphalidae). J Insect Behav 14:129–147. https://doi.org/10.1023/A:1007809915296 DOI

Kolar V, Boukal DS (2020) Habitat preferences of the endangered diving beetle Graphoderus bilineatus: implications for conservation management. Insect Conserv Divers 13:480–494. https://doi.org/10.1111/icad.12433 DOI

Konvicka M, Nedved O, Fric Z (2002) Early-spring floods decrease the survival of hibernating larvae of a wetland-inhabiting population of Neptis rivularis (Lepidoptera: Nymphalidae). Acta Zool Acad Sci Hung 48:79–88

Konvicka M, Fric Z, Benes J (2006) Butterfly extinctions in European states: do socioeconomic conditions matter more than physical geography? Global Ecol Biogeogr 15:82–92. https://doi.org/10.1111/j.1466-822X.2006.00188.x DOI

Konvickova H, Spitzer L, Fric ZF, Kepka P, Lestina D, Novotny D, Zapletal M, Zimmermann K, Maresova JP, Benes J, Konvicka M (2023) Perishing rich, expanding poor: demography and population genetic patterns in two congeneric butterflies. Mol Ecol 32:575–594. https://doi.org/10.1111/mec.16784 PubMed DOI

Kral K (2022) Flight behaviour of the woodland butterfly Neptis rivularis (Scopoli, 1763) (Nymphalidae) within residential gardens. Acta Entomol Slovenica 30:21–30

Krenn HW (2008) Feeding behaviours of neotropical butterflies (Lepidoptera, Papilionoidea). Denisia, Zugleich Kataloge Der Oberösterreichischen Landesmuseen Neue Serie 88: 295–304

Kudrna O (2002) The distribution atlas of European butterflies. Oedippus 20:1–343

Kudrna O (1970) Butterflies of South Bohemia. Entomol Rec J Var 82, 323–330 + 83, 53–67

Kudrna O (1986) Butterflies of Europe. Volume 8. Aspects of the conservation of butterflies in Europe. AULA Verlag, Wiesbaden

Lebeau J, Wesselingh RA, van Dyck H (2016) Nectar resource limitation affects butterfly flight performance and metabolism differently in intensive and extensive agricultural landscapes. Proc R Soc B Biol 283:20160455. https://doi.org/10.1098/rspb.2016.0455 DOI

Loos J, Schroer C, Becker T, Kastal A, Kortmann E, Dolek M (2021) Mosaic landscapes provide conservation pockets for an endangered species: Colias myrmidone in Romania. Insect Conserv Diversity 15:359–369. https://doi.org/10.1111/icad.12561 DOI

Lorimer J, Sandom C, Jepson P, Doughty C, Barua M, Kirby KJ (2015) Rewilding: science, practice, and politics. Ann Rev Environ Res 40:39–62. https://doi.org/10.1146/annurev-environ-102014-021406 DOI

Maes D, Van Calster H, Herremans M, Van Dyck H (2022) Challenges and bottlenecks for butterfly conservation in a highly anthropogenic region: Europe’s worst case scenario revisited. Biol Conserv 274:109732. https://doi.org/10.1016/j.biocon.2022.109732 DOI

Marchant NC, Purwanto A, Harsanto FA, Boyd NS, Harrison ME, Houlihan PR (2015) ‘Random-flight’ dispersal in tropical fruit-feeding butterflies? High mobility, long lifespans and no home ranges. Ecol Entomol 40:696–706. https://doi.org/10.1111/een.12239 DOI

Martelli F, Paradiso F, Ghidotti S, Viterbi R, Cerrato C, Bonelli S (2022) Invasion patterns and niche comparison of the butterfly Cacyreus marshalli among native and non-native range. Biol Invasions 24:3935–3952. https://doi.org/10.1007/s10530-022-02895-3 DOI

Middleton BA, Holsten B, Van Diggelen R (2006) Biodiversity management of fens and fen meadows by grazing, cutting and burning. Appl Veg Sci 9:307–316. https://doi.org/10.1111/j.1654-109X.2006.tb00680.x DOI

Moller H, Tilley JAV (1989) Beech honeydew – seasonal variation and use by wasps, honey bees, and other insects. N Z J Zool 16:289–302. https://doi.org/10.1080/03014223.1989.10422894 DOI

Musil P (1998) Changes in numbers of breeding populations of water birds of fishponds in the Třeboň Basin in the years 1981–1997. Sylvia 34:13–26

Niitepold K, Parry HA, Harris NR, Appel AG, de Roode JC, Kavazis AN, Hood WR (2022) Flying on empty: reduced mitochondrial function and flight capacity in food-deprived monarch butterflies. J Exp Biol 225:244431. https://doi.org/10.1242/jeb.244431 DOI

Novák I, Spitzer K (1982) Ohrožený svět hmyzu [Endangered world of insects]. Academia, Praha

Öckinger E, Eriksson AK, Smith HG (2006) Effects of grassland abandonment, restoration and management on butterflies and vascular plants. Biol Conserv 133:291–300. https://doi.org/10.1016/j.biocon.2006.06.009 DOI

Odendaal FJ, Iwasa Y, Ehrlich PR (1985) Duration of female availability and its effect on butterfly mating systems. Am Nat 125:673–678. https://doi.org/10.1086/284370 DOI

Polic D, Tamario C, Franzen M, Betzholtz PE, Yildirim Y, Forsman A (2021) Movements and occurrence in two closely related fritillary species. Ecol Entomol 46:428–439. https://doi.org/10.1111/een.12987 DOI

Prach K (1993) Vegetational changes in a wet meadow complex, South Bohemia, Czech Republic. Folia Geobot Phytotax 28:1–13. https://doi.org/10.1007/BF02853197 DOI

Prach K, Straškrabová J (1996) Restoration of degraded meadows: an experimental approach. In: Prach K, Jeník J, Large ARG (eds) Floodplain ecology and management, the Luznice River in the Trebon Biosphere Reserve, Central Europe. SPB Academic, Amsterdam, pp 87–93

Räuschl G (2002) Zu Ökologie und Artenschutz des Schwarzen Trauerfalters (Neptis rivularis SCOPOLI, 1763) (Lepidoptera: Nymphalidae) in Wien-Ottakring, Österreich. Beitr Entomofaunistik 3:81–85

Reinhardt R, Harpke A, Caspari S, Dolek M, Kühn E, Musche M, Trusch R, Wiemers W, Settele J (2020) Verbreitungsatlas der Tagfalter und Widderchen Deutschlands. Verlag Eugen Ulmer

Richardson ID (2019) Revision of the genus Neptis Fabricius, 1807 (Lepidoptera, Nymphalidae) in the Afrotropical Region: currently described taxa. Metamorphosis 30:69–221 DOI

Rosenzweig ML (2003) Win-win ecology: how the earth’s species can survive in the midst of human enterprise. Oxford University Press, Oxford DOI

Russi D, ten Brink P, Farmer A, Badura T, Coates D, Förster J, Kumar R, Davidson N (2013) The economics of ecosystems and biodiversity for water and wetlands. IEEP, London and Brussels; Ramsar Secretariat, Gland

Schmitt T, Habel JC, Besold J, Becker T, Johnen L, Knolle M, Rzepecki A, Schultze J, Zapp A (2006) The Chalk-hill Blue Polyommatus coridon (Lycaenidae, Lepidoptera) in a highly fragmented landscape: how sedentary is a sedentary butterfly? J Insect Conserv 10:311–316. https://doi.org/10.1007/s10841-006-9000-6 DOI

Schtickzelle N, Le Boulengé E, Baguette M (2002) Metapopulation dynamics of the bog fritillary butterfly: demographic processes in a patchy population. Oikos 97(3):349–360. https://doi.org/10.1034/j.1600-0706.2002.970305.x DOI

Scott JA (1974) Mate-locating behavior of butterflies. Am Midl Nat 91:103–117. https://doi.org/10.2307/2424514 DOI

Sielezniew M, Kostro-Ambroziak A, Klimczuk P, Deoniziak K, Pałka K, Nowicki P (2019) Habitat-related differences in the adult longevity of two ecotypes of a specialized butterfly. J Zool 307:93–103. https://doi.org/10.1111/jzo.12625 DOI

Sielezniew M, Kostro-Ambroziak A, Korosi A (2020) Sexual differences in age-dependent survival and life span of adults in a natural butterfly population. Sci Rep 10:10394. https://doi.org/10.1038/s41598-020-66922-w PubMed DOI PMC

Sietses DJ, Faupin G, de Boer WF, de Jong CB, Henkens RJHG, Usukhjargal D, Batbaatar T (2009) Resource partitioning between large herbivores in Hustai National Park, Mongolia. Mammal Biol 74:381–393. https://doi.org/10.1016/j.mambio.2009.04.001 DOI

Skelly DK, Halverson MA, Freidenburg LK, Urban MC (2005) Canopy closure and amphibian diversity in forested wetlands. Wetl Ecol Manag 13:261–268. https://doi.org/10.1007/s11273-004-7520-y DOI

Slamova I, Klecka J, Konvicka M (2011) Diurnal behavior and habitat preferences of Erebia aethiops, an aberrant lowland species of a mountain butterfly clade. J Insect Behav 24:230–246. https://doi.org/10.1007/s10905-010-9250-8 DOI

Slamova I, Klecka J, Konvicka M (2013) Woodland and grassland mosaic from a butterfly perspective: habitat use by Erebia aethiops (L epidoptera: S atyridae). Insect Conserv Divers 6:243–254.  https://doi.org/10.1111/j.1752-4598.2012.00212.x

Spitzer K (1958) K výskytu a bionimii Neptis coenobita Stoll. subsp. innominata Lewis v jižních Čechách (Lep., Nymphalidae). Čas Čs Spol Entomol 55:298–299

Spitzer L, Beneš J (2022) Expansion of the Marbled Fritillary (Brenthis daphne) (Lepidoptera: Nymphalidae) in Moravia and Silesia. Acta Carpathica Occidentalis 13:68–76 DOI

Spitzer K, Bezděk A, Jaroš J (1999) Ecological succession of a relict Central European peat bog and variability of its insect biodiversity. J Insect Conserv 3:97–106. https://doi.org/10.1023/A:1009634611130 DOI

Stephens PA, Houston AI, Harding KC, Boyd IL, McNamara JM (2014) Capital and income breeding: the role of food supply. Ecology 95:882–896. https://doi.org/10.1890/13-1434.1 PubMed DOI

Strausz M, Fiedler K, Franzén M, Wiemers M (2012) Habitat and host plant use of the large copper butterfly Lycaena dispar in an urban environment. J Insect Conserv 16:709–721. https://doi.org/10.1007/s10841-012-9456-5 DOI

Sucháčková Bartoňová A, Konvička M, Marešová J, Bláhová D, Číp D, Skala P, Andres M, Hula V, Dolek M, Geyer A, Böck O, Kadlec T, Fric ZF (2021) Extremely endangered butterflies of scattered central European dry grasslands under current habitat alteration. Insect Systematics Divers 5:6. https://doi.org/10.1093/isd/ixab017 DOI

Tammaru T, Haukioja E (1996) Capital breeders and income breeders among Lepidoptera: consequences to population dynamics. Oikos 77:561–564. https://doi.org/10.2307/3545946 DOI

Thomas JS (1983) The ecology and conservation of Lysandra bellargus (Lepidoptera, Lycaenidae) in Britain. J Appl Ecol 20:59–83. https://doi.org/10.2307/2403376 DOI

Tseng H-Y, Chiba H, Lohman DJ, Yen S-H, Aduse-Poku K, Ohshima Y, Wu L-W (2022) Out of Asia: intercontinental dispersals after the Eocene-Oligocene transition shaped the zoogeography of Limenitidinae butterflies (Lepidoptera: Nymphalidae). Mol Phyl Evol 170:107444. https://doi.org/10.1016/j.ympev.2022.107444 DOI

Vera FWM (2000) Grazing ecology and forest history. CABI, Wallingford DOI

Vetter SG, Ruf T, Bieber C, Arnold W (2015) What is a mild winter? Regional differences in within-species responses to climate change. PLS One 10:e0132178. https://doi.org/10.1371/journal.pone.0132178 DOI

Vinther E, Hald AB (2000) Restoration of an abandoned species-rich fen-meadow in Denmark changes in species richness and dynamics of plant groups during 12 years. Nord J Bot 20:573–584. https://doi.org/10.1111/j.1756-1051.2000.tb01608.x DOI

Vlasanek P, Hauck D, Konvicka M (2009) Adult sex ratio in the Parnassius mnemosyne butterfly: effects of survival, migration, and weather. Isr J Ecol Evol 55:233–252. https://doi.org/10.1560/IJEE.55.3.233 DOI

Vlasanek P, Fric ZF, Zimmermann K, Novotný D, Čížek O, Klečková I, Vrba P, Kadlec T, Konvička M (2018) Do butterfly activity data from mark-recapture surveys reflect temporal patterns? J Insect Behav 31:385–401. https://doi.org/10.1007/s10905-018-9686-9 DOI

Wagner DL (2020) Insect declines in the Anthropocene. Ann Rev Entomol 65:457–480. https://doi.org/10.1146/annurev-ento-011019-025151 DOI

Warren MS, Maes D, van Swaay CA, Goffart P, Van Dyck H, Bourn NA, Wynhoff I, Hoare D, Ellis S (2021) The decline of butterflies in Europe: problems, significance, and possible solutions. Proc Nat Acad Sci USA 118:2002551117. https://doi.org/10.1073/pnas.2002551117 DOI

Weyer J, Schmitt T (2013) Knowing the way home: strong philopatry of a highly mobile insect species, Brenthis ino. J Insect Conserv 17:1197–1208. https://doi.org/10.1007/s10841-013-9601-9 DOI

Wnorowska K (2021) Monitoring proposal of the species Neptis rivularis (Scopoli, 1763) in Poland. Folia Pomeranae Univ Technol Stetin 361:35–49

Zimmermann K, Fric Z, Filipova L, Konvicka M (2005) Adult demography, dispersal and behaviour of Brenthis ino (Lepidoptera: Nymphalidae): how to be a successful wetland butterfly. Eur J Entomol 102:699–706. https://doi.org/10.14411/eje.2005.100 DOI