Eur. J. Entomol. 123: 122-133, 2026 | DOI: 10.14411/eje.2026.012

Influence of extreme climatic effects on butterfly assemblages in a Mediterranean landscape, with new records of threatened and endemic speciesOriginal article

Juan José LUCAS LÓPEZ ORCID...1, Juan Ignacio ARCE CRESPO ORCID...1, 2, *, José María REY-BENAYAS ORCID...3, 4
1 Entomological and Environmental Society of Castilla-La Mancha (SEACAM), C/ Olivar, 4, 1ºA, 45593 Bargas (Toledo), Spain; e-mail: juanjoseminerales1995@gmail.com
2 Society for the Conservation and Study of Butterflies in Spain (SOCEME), C/ Darwin, 2, Fuencarral-El Pardo, 28049 Madrid, Spain; e-mail: jiarce.bio@gmail.com
3 Department of Life Sciences - Ecology and Forest Restoration Group (FORECO), Science Building, University of Alcalá, 28805 Alcalá de Henares, Madrid, Spain; e-mail: josem.rey@uah.es
4 Fundación Internacional para la Restauración de Ecosistemas, C/ Princesa, 3 dpdo., Apto. 703, 28008 Madrid, Spain

Climate change is seriously threatening biodiversity and, especially, insects because their biological cycles are highly dependent on environmental conditions. We studied (1) how climate characterised by droughts and heatwaves affected the butterfly assemblage at four sampled sites in Campo de Montiel (central Spain) over three consecutive years; and (2) the differences in the assemblage abundance, species richness, density, and diversity among sites that represent a variety of land use/cover types. Sites were sampled with transects using the Butterfly Monitoring Scheme methodology (Pollard & Yates, 1993). A total of 8,275 butterflies and 69 species were sampled across sites and years. Sites were well sampled in the three years according to species richness estimators and accumulation curves. The relationship between the mean length of dry spells and butterfly abundance was negative and significant at all sites. However, heatwaves did not affect butterfly abundance and species richness. There were no significant differences in the ecological parameters studied among sites. Mediterranean forest transects tended to have higher values of ecological parameters than those dominated by more degraded vegetation, pine plantations and cropland intensification. Butterfly assemblage and diversity were positively related to the presence of semi-natural habitat with traditional agricultural and livestock uses. Our surveys recorded Polyommatus celina for the first time in Castilla-La Mancha. Two endangered and endemic species, Polyommatus violetae and Kretania hesperica, were recorded at the site with higher Mediterranean forest cover, which increases the conservation value of this site. Maintaining semi-natural habitats and traditional farming systems may allow butterfly assemblages to be more resilient to climate change.

Keywords: Lepidoptera, conservation, Butterfly Monitoring Scheme, climate change, drought, heatwave

Received: July 17, 2025; Revised: February 17, 2026; Accepted: February 17, 2026; Published online: March 31, 2026  Show citation

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LUCAS LÓPEZ, J.J., ARCE CRESPO, J.I., & REY-BENAYAS, J.M. (2026). Influence of extreme climatic effects on butterfly assemblages in a Mediterranean landscape, with new records of threatened and endemic species. EJE, 123, Article 122-133. https://doi.org/10.14411/eje.2026.012
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    References

    1. Abarca M., Larsen E.A. & Ries L. 2019: Heatwaves and novel host consumption increase overwinter mortality of an imperiled wetland butterfly. - Front. Ecol. Evol 7: 193, 13 pp. Go to original source...
    2. Alfaro F.M. & Pizarro-Araya J. 2017: Estimación de la riqueza de coleópteros epigeos de la Reserva Nacional Pingüino de Humboldt (Regiones de Atacama y Coquimbo, Chile). - Gayana 81: 39-51. Go to original source...
    3. Ali E., Cramer W., Carnicer J., Georgopoulou E., Hilmi N.J.M., Le Cozannet G. & Lionello P. 2022: Cross-Chapter Paper 4: Mediterranean Region. In Pörtner H.O., Roberts D.C., Tignor M., Poloczanska E.S., Mintenbeck K., Alegrîa A., Craig M., Langsdorf S., Löschke S., Möller V., Okem A. & Rama B. (eds): Climate Change 2022: Impacts, Adaptation and Vulnerability. Contribution of Working Group II to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, UK and New York, NY, pp. 2233-2272. Go to original source...
    4. Andreas E. & Mevi-Schütz J. 2009: Adult food resources in butterflies. In Settele J., Shreeve T., Konvicka M. & Van Dyck H. (eds): Ecology of Butterflies in Europe. Cambridge University Press, Cambridge, UK, pp. 9-16.
    5. Araújo M.B. & Guisan A. 2006: Five (or so) challenges for species distribution modelling. - J. Biogeogr. 33: 1677-1688. Go to original source...
    6. Arce J.I. 2024: Biología, distribución y conservación de los lepidópteros diurnos amenazados de Castilla-La Mancha. PhD thesis, Universidad de Castilla-La Mancha, Toledo, xii + 260 pp.
    7. Balmaki B., Rostami M.A., Allen J.M. & Dyer L.A. 2024: Effects of climate change on Lepidoptera pollen loads and their pollination services in space and time. - Oecologia 204: 751-759. Go to original source...
    8. Bartomeus I., Ascher J.S., Wagner D., Danforth B.N., Colla S., Kornbluth S. & Winfree R. 2011: Climate-associated phenological advances in bee pollinators and bee-pollinated plants. - Proc. Natl. Acad. Sci. U.S.A. 108: 20645-20649. Go to original source...
    9. Bautista-Hernández C.E., Monks S. & Pulido-Flores G. 2013: Los parásitos y el estudio de su biodiversidad: un enfoque sobre los estimadores de la riqueza de especies. Estudios científicos en el estado de Hidalgo y zonas aledañas. 4. https://digitalcommons.unl.edu/hidalgo/4.
    10. Baz A. 2002: Nectar plant sources for the threatened Apollo butterfly (Parnassius apollo L. 1758) in populations of central Spain. - Biol. Conserv. 103: 277-282. Go to original source...
    11. Bonelli S., Cerrato C., Barbero F., Boiani M.V., Buffa G., Casacci L.P., Fracastoro L., Provenzale A., Rivella E., Zaccagno M. & Balleto E. 2022: Changes in alpine butterfly communities during the last 40 years. - Insects 13: 43, 22 pp. Go to original source...
    12. Chao A. 1984: Non-parametric estimation of the number of classes in a population. - Scand. J. Stat. 11: 265-270.
    13. Chao A. 1987: Estimating the population size for capture-recapture data with unequal catchability. - Biometrics 43: 783-791. Go to original source...
    14. Chao A., Hwang W.H., Chen Y.C. & Kuo C.Y. 2000: Estimating the number of shared species in two communities. - Stat. Sin. 10: 227-246.
    15. Chazdon R.L., Colwell R.K., Denslow J.S. & Guariguata M.R. 1998: Statistical methods for estimating species richness of woody regeneration in primary and secondary rain forests of NE Costa Rica. In Dallmeier F. & Comiskey J.A. (eds): Forest Biodiversity Research, Monitoring and Modeling: Conceptual Background and Old World Case Studies. Parthenon Publishing, Paris, pp. 285-309. Go to original source...
    16. Cleary D.F.R. 2004: Assessing the use of butterflies as indicators of logging in Borneo at three taxonomic levels. - J. Econ. Entomol 97: 429-435. Go to original source...
    17. Colwell R.K. 2013: EstimateS (Version 9.1.0). URL: http://purl.oclc.org/estimates.
    18. Corbet S.A. 2000: Butterfly nectaring flowers: butterfly morphology and flower form. - Entomol. Exp. Appl. 96: 289-298. Go to original source...
    19. Edwards C.B., Zipkin E.F., Henry E.H., Haddad N.M., Forster M.L., Burls K.J., Campbell S.P., Crone E.E., Diffendorfer J., Douglas M.R. et al. 2025: Rapid butterfly declines across the United States during the 21st century. - Science 387: 1090-1094. Go to original source...
    20. Fernández-Rubio F. 1976: Genitalias (andropigios) de los ropalóceros de Álava y su entorno ibérico. Parte I: Lycaenidae. Diputación Foral de Álava, Vitoria, 80 pp.
    21. Fernández-Rubio F. 1977: Genitalias (andropigios) de los ropalóceros de Álava y su entorno ibérico. Parte II: Libytheidae, Nymphalidae, Danaidae. Diputación Foral de Álava, Vitoria, 54 pp.
    22. Fernández-Rubio F. 1981: Genitalias (andropigios) de los ropalóceros de Álava y su entorno ibérico. Parte III: Nemeobidae, Pieridae, Papilionidae, Hesperidae. Diputación Foral de Álava, Vitoria, 65 pp.
    23. Fernández-Rubio F. 1982: Genitalias (andropigios) de los ropalóceros de Álava y su entorno ibérico. Parte IV: Satyridae. Diputación Foral de Álava, Vitoria, 58 pp.
    24. Forister M.L., Pelton E.M. & Black S.H. 2019: Declines in insect abundance and diversity: we know enough to act now. - Conserv. Sci. Pract. 1: e80, 8 pp. Go to original source...
    25. Forister M.L, Grames E.M., Halsch C.A., Burls K.J., Carroll C.F., Bell K.L., Jahner J.P., Bradford T.A., Zhang J., Cong Q. et al 2023: Assessing risk for butterflies in the context of climate change, demographic uncertainty, and hetero­genous data sources. - Entomol. Monogr. 93: e1584, 25 pp. Go to original source...
    26. Frank D., Reichstein M., Bahn M., Thonicke K. Frank D., Mahecha M.D., Smith P., van der Velde M., Vicca S., Babst F. et al. 2015: Effects of climate extremes on the terrestrial carbon cycle: Concepts, processes and potential future impacts. - Glob. Chang. Biol. 21: 2861-2880. Go to original source...
    27. Frigero M.L.P., Boaro C.S.F., Galetto L., Tunes P. & Guimarães E. 2025: Extreme events induced by climate change alter nectar offer to pollinators in cross pollination-dependent crops. - Sci. Rep. 15: 10852, 12 pp. Go to original source...
    28. Garcîa-Barros E., Munguira M.L., Stefanescu C. & Vives Moreno A. 2013: Lepidoptera, Papilionoidea. In Ramos M.A. et al. (eds): Fauna Ibérica, Vol. 37. Museo Nacional de Ciencias Naturales, CSIC, Madrid, 1213 pp.
    29. Gordo O. & Sanz J.J. 2005: Phenology and climate change: a long-term study in a Mediterranean locality. - Oecologia 146: 484-495. Go to original source...
    30. Habel J.C., Schmitt T., Gros P. & Ulrich W. 2022: Breakpoints in butterfly decline in Central Europe over the last century. - Sci. Total Environ. 851: 158315, 9 pp. Go to original source...
    31. Hallmann C.A., Sorg M., Jongejans E., Siepel H., Hofland N., Schwan H., Stenmans W., Müller A., Sumser H., Hörren T. et al. 2017: More than 75 percent decline over 27 years in total flying insect biomass in protected areas. - PLoS ONE 12: e0185809, 21 pp. Go to original source...
    32. Halsch C.A., Shapiro A.M., Thorne J.H., Rodman K.C., Parra A., Dyer L.A., Gompert Z., Smilanich A.M & Forister M.L. 2024: Thirty-six years of butterfly monitoring, snow cover, and plant productivity reveal negative impacts of warmer winters and increased productivity on montane species. - Glob. Change Biol. 30: e17044, 15 pp. Go to original source...
    33. Hansen J., Sato M. & Ruedy R. 2012: Perception of climate change. - Proc. Natl. Acad. Sci. U.S.A 109: 2415-2423. Go to original source...
    34. Hayes M.P. & Turner E.C. 2023: The impact of an extreme weather event on butterfly community activity, use of shelter and shade. Dataset. https://doi.org/10.6084/m9.figshare.22083029.v2. Go to original source...
    35. Hayes M.P., Ashe-Jepson E., Hitchcock G.E., Clark R., Hellon J., Knock R.I., Bladon A.J. & Turner E.C. 2024: Heatwave predicts a shady future for insects: impacts of an extreme weather event on a chalk grassland in Bedfordshire, UK. - J. Insect Conserv. 28: 923-933. Go to original source...
    36. IBM Corp. Released 2020: IBM SPSS Statistics for Windows, Version 27.0. URL: https://www.ibm.com/docs/en/spss-statistics/27.0.0.
    37. IPCC 2014: Climate change 2014: Impacts, adaptation, and vulnerability. In Core Writing Team, Pachauri R.K. & Meyer L.A. (eds): Contribution of Working Groups I, II and III to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. IPCC, Geneva, 151 pp.
    38. IPCC 2023: Climate change 2023: Synthesis report. In Core Writing Team, Lee H. & Romero J. (eds): Contribution of Working Groups I, II and III to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. IPCC, Geneva, pp. 35-115.
    39. Jaworski C.C., Geslin B., Zakardjian M., Lecareux C., Caillault P., Nève G., Meunier J.-Y., Dupouyet S., Sweeney A.C.T., Lewis O.T., Dicks L.V. & Fernandez C. 2022: Long-term experimental drought alters floral scent and pollinator visits in a Mediterranean plant community despite overall limited impacts on plant phenotype and reproduction. - J. Ecol. 110: 2628-2648. Go to original source...
    40. Jiménez-Valverde A., Martîn Cano J. & Munguira M.L. 2004: Patrones de diversidad de la fauna de mariposas del Parque Nacional de Cabañeros y su entorno (Ciudad Real, España central) (Lepidoptera, Papilionoidea, Hesperioidea). - Anim. Biodiv. Conserv. 27: 15-24. Go to original source...
    41. Johansson V., Kindvall O., Askling J., Säwenfalk D.S., Norman H. & Franzén M. 2022: Quick recovery of a threatened butterfly in well-connected patches following an extreme drought. - Insect Conserv. Divers. 15: 572-582. Go to original source...
    42. Kassambara A. 2023: ggpubr: 'ggplot2' Based Publication Ready Plots. R Package Version 0.6.0. URL: https://rpkgs.datanovia.com/ggpubr/.
    43. Kitahara M., Yumoto M. & Kobayashi T. 2008: Relationship of butterfly diversity with nectar plant species richness in and around the Aokigahara primary woodland of Mount Fuji, central Japan. - Biodiv. Conserv. 17: 2713-2734. Go to original source...
    44. Klein Tank A.M.G., Zwiers F.W. & Zhang X. 2009: Guidelines on analysis of extremes in a changing climate in support of informed decisions for adaptation. - Climate Data and Monitoring WCDMP No. 72, 52 pp.
    45. Klockmann M. & Fischer K. 2017: Effects of temperature and drought on early life stages in three species of butterflies: mortality of early life stages as a key determinant of vulnerability to climate change? - Ecol. Evol. 7: 10871-10879. Go to original source...
    46. Kremen C. 1992: Assessing the indicator properties of species assemblages for natural areas monitoring. - Ecol. Appl 2: 203-217. Go to original source...
    47. Lister B.C. & Garcia A. 2018: Climate-driven declines in arthropod abundance restructure a rainforest food web. - Proc. Natl. Acad. Sci. U.S.A. 115: 10397-10406. Go to original source...
    48. Maes D., Verovnik R., Wiemers M., Brosens D., Beshkov S., Bonelli S., Buszko J., Cantú-Salazar L., Cassar L.F., Collins S. et al. 2019: Integrating national Red Lists for prioritising conservation actions for European butterflies. - J. Insect Conserv. 23: 301-330. Go to original source...
    49. Meehl G.A. & Tebaldi C. 2004: More intense, more frequent, and longer lasting heat waves in the 21st century. - Science 305: 994-997. Go to original source...
    50. Melone G.G., Stuligross C. & Williams N.M. 2024: Heatwaves increase larval mortality and delay development of a solitary bee. - Ecol. Entomol 49: 433-444. Go to original source...
    51. Microsoft Corporation 2018: Microsoft Excel. URL: https://office.microsoft.com/excel.
    52. Mingarro M., Cancela J.P., Burón-Ugarte A., Garcîa-Barros E., Munguira M.L., Romo H. & Wilson R.J. 2021: Butterfly communities track climatic variation over space but not time in the Iberian Peninsula. - Insect Conserv. Divers. 14: 647-670. Go to original source...
    53. Montagud S. & Garcîa-Alamá J.A. 2010: Mariposas diurnas de la Comunitat Valenciana (Papilionoidea and Hesperioidea). Colección Biodiversidad 17, Conselleria de Medi Ambient, Aigua, Urbanisme i Habitatge, Generalitat Valenciana, Valencia, 472 pp.
    54. Mora A., Wilby A. & Menéndez R. 2022: Abandonment of cultural landscapes: butterfly communities track the advance of forest over grasslands. - J. Insect Conserv. 26: 85-96. Go to original source...
    55. Munguira M.L., Barea-Azcón J.M., Castro S., Olivares J. & Miteva S. 2015: Species Recovery Plan for the Andalusian Anomalous Blue (Polyommatus violetae). Butterfly Conservation Europe, Wageningen, 30 pp.
    56. Munguira M.L., Barea-Azcón J.M., Castro-Cobo S., Garcîa-Barros E., Miteva S., Olivares J. & Romo H. 2017: Ecology and recovery plans for the four Spanish endangered endemic butterfly species. - J. Insect Conserv. 21: 423-437. Go to original source...
    57. Ovaskainen O., Skorokhodova S., Yakovleva M., Sukhov A., Kutenkov A., Kutenkova N., Shcherbakov A., Meyke E. & Delgado M.M. 2013: Community-level phenological response to climate change. - Proc. Natl. Acad. Sci. U.S.A. 110: 13434-13439. Go to original source...
    58. Parmesan C. 2006: Ecological and evolutionary responses to recent climate change. - Annu. Rev. Ecol. Evol. Syst. 37: 637-669. Go to original source...
    59. Parmesan C. 2007: Influences of species, latitudes and methodologies on estimates of phenological response to global warming. - Glob. Change Biol. 13: 1860-1872. Go to original source...
    60. Parmesan C. & Yohe G. 2003: A globally coherent fingerprint of climate change impacts across natural systems. - Nature 421: 37-42. Go to original source...
    61. Parmesan C., Ryrholm N., Stefanescu C., Hill J.K., Thomas C.D., Descimon H., Huntley B., Kaila L., Kullberg J., Tammaru T. et al. 1999: Poleward shifts in geographical ranges of butterfly species associated with regional warming. - Nature 399: 579-583. Go to original source...
    62. Peñuelas J., Filella I. & Comas P. 2002: Changed plant and animal life cycles from 1952 to 2000 in the Mediterranean region. - Glob. Change Biol. 8: 531-544. Go to original source...
    63. Pérez-Fernández R. 2011: Plebejus hespericus (Rambur, 1840) en el centro de la Península Ibérica. Distribución geográfica, caracterización del hábitat, parasitoides y conservación (Lepidoptera: Lycaenidae). - Shilap Rev. Lepidopt. 39: 325-344.
    64. Phillips B.B., Shaw R.F., Holland M.J., Fry E.L., Bardgett R.D., Bullock J.M. & Osborne J.L. 2018: Drought reduces floral resources for pollinators. - Glob. Change Biol. 24: 3226-3235. Go to original source...
    65. Pinheiro J.C. & Bates D.M. 2000: Mixed-Effects Models in S and S-PLUS. Springer, New York, 528 pp. Go to original source...
    66. Pinheiro J.C. & Bates D.M. 2023: nlme: Linear and Nonlinear Mixed Effects Models. R Package Version 3.1-164. R Core Team. URL: https://CRAN.R-project.org/package=nlme.
    67. Pla L. & Matteucci S.D. 2001: Bootstrap confidence intervals for Shannon biodiversity index. - Revta Fac. Agron. Univ. Zulia 18: 222-234.
    68. Pollard E. & Yates T.J. 1993: Monitoring Butterflies for Ecology and Conservation. The British Butterfly Monitoring Scheme. Springer, Dordrecht, 292 pp.
    69. QGIS Association 2021: QGIS Geographic Information System. QGIS Association. URL: http://www.QGIS.org.
    70. R Core Team 2021: R: A Language and Environment for Statistical Computing. Vienna. URL: https://www.r-project.org/.
    71. Radchuk V., Turlure C. & Schtickzelle N. 2013: Each life stage matters: the importance of assessing the response to climate change over the complete life cycle in butterflies. - J. Anim. Ecol. 82: 275-285. Go to original source...
    72. Romo H., Garcîa-Barros E., Wilson R.J., Mateo R.G. & Munguira M.L. 2023: Modelling the scope to conserve an endemic-rich mountain butterfly taxon in a changing climate. - Insect Conserv. Divers. 16: 451-467. Go to original source...
    73. Sevilleja C.G., Van Swaay C.A.M., Bourn N., Collins S., Settele J., Warren M.S., Wynhoff I. & Roy D.B. 2019: Butterfly Transect Counts: Manual to Monitor Butterflies. Report VS2019.016, Butterfly Conservation Europe and De Vlinderstichting/Dutch Butterfly Conservation, Wageningen, 16 pp.
    74. Shannon C.E. & Weaver W. 1949: The Mathematical Theory of Communication. University of Illinois, Urbana, IL, 144 pp.
    75. Shirey V., Neupane N., Guralnick R. & Ries L. 2024: Rising minimum temperatures contribute to 50 years of occupancy decline among cold-adapted Arctic and boreal butterflies in North America. - Glob. Change Biol. 30: e17205, 16 pp. Go to original source...
    76. Smith E.P. & Van Belle G. 1984: Nonparametric estimation of species richness. - Biometrics 40: 119-129. Go to original source...
    77. Stefanescu C., Torre I., Jubany J. & Páramo F. 2011: Recent trends in butterfly populations from north-east Spain and Andorra in the light of habitat and climate change. - J. Insect Conserv. 15: 83-93. Go to original source...
    78. Thomas J.A., Telfer M.G., Roy D.B., Preston C.D., Greenwood J.J.D., Asher J., Fox R., Clarke R.T. & Lawton J.H. 2004: Comparative losses of British butterflies, birds, and plants and the global extinction crisis. - Science 303: 1879-1881. Go to original source...
    79. Tolman T. & Lewington R. 2002: Guîa de campo de las mariposas de España y de Europa. Lynx Editions, Barcelona, 320 pp.
    80. Ubach A., Páramo F., Prohom M. & Stefanescu C. 2022: Weather and butterfly responses: a framework for understanding population dynamics in terms of species' life-cycles and extreme climatic events. - Oecologia 199: 427-439. Go to original source...
    81. Ummenhofer C.C. & Meehl G.A. 2017: Extreme weather and climate events with ecological relevance: a review. - Phil. Trans. R. Soc. (B) 372: 20160135, 13 pp. Go to original source...
    82. Van Dyck H., Puls R., Bonte D., Gotthard K. & Maes D. 2015: The lost generation hypothesis: could climate change drive ectotherms into a developmental trap? - Oikos 124: 54-61. Go to original source...
    83. Van Halder I., Barbaro L., Corcket E. & Jactel H. 2008: Importance of seminatural habitats for the conservation of butterfly communities in landscapes dominated by pine plantations. - Biodiv. Conserv. 17: 1149-1169. Go to original source...
    84. Van Klink R., Bowler D.E., Gongalsky K.B., Shen M., Swengel S.R. & Chase J.M. 2024: Disproportionate declines of formerly abundant species underlie insect loss. - Nature 628: 359-364. Go to original source...
    85. Van Swaay C.A.M. & Warren M.S. 2012: Developing Butterflies as Indicators in Europe: Current Situation and Future Options. De Vlinderstichting/Dutch Butterfly Conservation, Butterfly Conservation UK, Butterfly Conservation Europe, Wageningen, report no. VS2012.012, 24 pp.
    86. Van Swaay C.A.M., Maes D. & Warren M.S. 2009: Conservation status of European butterflies. In Settele J., Shreeve T., Konvièka M. & Van Dyck H. (eds): Ecology of Butterflies in Europe. Cambridge University Press, Cambridge, pp. 322-338.
    87. Van Swaay C.A.M, Cuttelod A., Collins S., Maes D., López Munguira M., ©a¹iæ M., Settele J., Verovnik R., Verstrael T., Warre M. et al. 2010: European Red List of Butterfies. Publications Office of the European Union, Luxembourg, 47 pp.
    88. Van Swaay C.A.M., Warren M.S., Ellis S., Clay J., Bellotto V., Allen D.J. & Trottet A. 2025a: Measuring the Pulse of European Biodiversity Using the Red List. European Red List of Butterflies. European Commission, Brussels, 80 pp.
    89. Van Swaay C., Ellis S. & Warren M. 2025b: Kretania hesperica. URL: https://dx.doi.org/10.2305/IUCN.UK.2025-1.RLTS.T39489A211108195.en. Go to original source...
    90. Van Swaay C., Ellis S. & Warren M. 2025c: Polyommatus violetae. URL: https://dx.doi.org/10.2305/IUCN.UK.2025-1.RLTS.T275427561A211448671.en. Go to original source...
    91. Verdú J.R., Numa C. & Galante E. 2011: Atlas y libro rojo de los invertebrados amenazados de España (especies vulnerables), Vol. 1. Dirección General de Medio Natural y Política Forestal, Ministerio de Medio Ambiente, Medio Rural y Marino, Madrid, 595 pp.
    92. Wagner D.L., Fox R., Salcido D.M. & Dyer L.A. 2021: A window to the world of global insect declines: moth biodiversity trends are complex and heterogeneous. - Proc. Natl. Acad. Sci. U.S.A. 118: e2002549117, 8 pp. Go to original source...
    93. Warren M.S., Maes D., Van Swaay C.A.M., Goffart P., Van Dyck H., Bourn N.A.D., Wynhoff I., Hoare D. & Ellis S. 2021: The decline of butterflies in Europe: problems, significance, and possible solutions. - Proc. Natl. Acad. Sci. U.S.A. 118: e2002551117, 10 pp. Go to original source...
    94. Wickham H., Averick M., Bryan J., Chang W., D'Agostino McGowan L., François R., Grolemund G., Hayes A., Henry L., Hester J. et al. 2019: Welcome to the tidyverse. - J. Open Source Softw 4: 1686, 6 pp. Go to original source...
    95. Wiemers M., Balletto E., Dincă V., Fric Z.F., Lamas G., Lukhtanov V., Munguira M.L., van Swaay C.A.M, Vila R., Vliegenthart A. et al. 2018: An updated checklist of the European butterflies (Lepidoptera, Papilionoidea). - ZooKeys 811: 9-45. Go to original source...
    96. Wilson R.J. & Fox R. 2021: Insect responses to global change offer sign-posts for biodiversity and conservation. - Ecol. Entomol. 46: 699-717. Go to original source...
    97. Wilson R.J., Gutiérrez D., Gutiérrez J., Martínez D., Agudo R. & Monserrat V.J. 2005: Changes to the elevational limits and extent of species ranges associated with climate change. - Ecol. Lett. 8: 1138-1146. Go to original source...
    98. Wilson R.J., Bennie J., Lawson C.R., Pearson D., Ortuzar-Ugarte G. & Gutiérrez D. 2015: Population turnover, habitat use and microclimate at the contracting range margin of a butterfly. - J. Insect Conserv. 19: 205-216. Go to original source...
    99. Yang L.H. & Rudolf V.H.W. 2010: Phenology, ontogeny and the effects of climate change on the timing of species interactions. - Ecol. Lett. 13: 1-10. Go to original source...
    100. Zografou K., Kati V., Grill A., Wilson R.J., Tzirkalli E., Pamperis L.N. & Halley J.M. 2014: Signals of climate change in butterfly communities in a Mediterranean protected area. - PLoS ONE 9: e87245, 9 pp. Go to original source...
    101. Zuur A.F., Ieno E.N., Walker N.J., Saveliev A.A. & Smith G. 2009: Mixed Effects Models and Extensions in Ecology with R. Springer, New York. URL: http://dx.doi.org/10.1007/978-0-387-87458-6. Go to original source...

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