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Butterflies of European islands: the implications of the geography and ecology of rarity and endemicity for conservation

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Abstract

Depending on their faunal content islands can function as important ‘vehicles’ for conservation. In this study, we examine data on 440 butterfly species over 564 European islands in 10 island groups. To determine the status of the butterfly fauna, we have adopted two approaches, island-focused and species-focused, examined using principal components analysis and regression modelling. In the former, we relate species richness, rarity and endemicity to island geography (area, elevation, isolation and location in latitude and longitude); in the latter, species occurrence on islands is examined in relation to distribution, range, range boundaries, and altitudinal limits on the continent as well as species’ ecology (number of host plants) and morphology (wing expanse). Species on islands are also assessed for their status on the continental mainland, their distributional dynamics (extinctions, distribution changes) and conservation status (Red Data Book, European Habitat Directive, Species of European Conservation Concern and Bern Convention listing. Unexpectedly, we find that a large fraction of the European butterfly species is found on the islands (63.4%; 59% on small islands) comprising some 6.2% of the land area of Europe. Although species occurring on the islands tend, on the whole, to have lower conservation status and are not declining over Europe, 45 species are endemics restricted to the islands. Species richness shows only a weak locational pattern and is related as expected to isolation from the continental source and island area; but, both rarity and endemicity have distinctive geographical bias to southern Europe, on islands now under increasing pressure from climate change and increasingly intensive human exploitation. The vulnerability of species on islands is emphasised in the relationship of island occurrence (% occurrence and presence/absence of species on any island) with continental distributions. A large proportion of the variation (84%) is accounted by continental distribution, the southern range limit and lower altitudinal limit. Most species (69%) occur on very few islands (<5%). In view of ongoing species dynamics on islands, migrations and extinctions of species, island repositories of species depend in large part on conservation of butterflies at continental sources. The unique faunas and rare species on islands also depend on appropriate concern being given to the island faunas. Conservation of European islands is thus a two-way process, sustaining sources and conserving island refuges. Residuals from the regressions (islands with more or fewer species, rare and endemic species; species occurring more or less frequently than expected on islands) provide warning signals of regions and islands deserving immediate attention.

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Acknowledgements

We thank the following for providing data: Michael Archer [Hebrides], I. Beavis [Isles of Scilly], Eliseo H Fernández-Vidal [Balearics], Lin Gander [Cardigan Island, UK], Martin Gascoigne-Pees [Simi, Canary islands], Nick Greatorex-Davies [Flatholm, UK], Jean-Alain Guilloton [French islands], Herve Guyot [Corsica], Martin Honey [Balearics], L. Huldén [Finnish islands], M. Hull [Herm, Sark], Stephen Jeffcoate (Isle of Man), Gérard Luquet [Noirmoutier, France], Tony Nagypal [Norwegian islands], Alain Olivier and Rob Parker [Greek islands], Christian Perrein and Atlas entomologique régional (Nantes) [French islands], Ian Rippey [Irish islands], Paul Sammut [Malta], W. Sandison [Wales: Llanddwyn Island], Constantí Stefanescu [Spanish islands], Mike Taylor [Chios], Andrew Wakeham-Dawson [Macaronesia] and David Withrington [Croatia]. Our grateful thanks to a team from Sweden for providing data for the study: Leif Aarvik, Jan-Olov Björklund, Per-Eric Betzholz, Fredrik Sjöberg, Jan Uddén, and Göran Palmqvist. Our grateful thanks to Tim H. Sparks (CEH, Monks Wood, Cambridgeshire UK) for his valuable advice on statistical matters and to Martin Wiemers for a preliminary view to one of us (OK) from his own research on Coenonympha elbana. Thanks to referees for useful comments made and to Tim New for his kindly and professional guidance in piloting this product to the terminus.

Author information

Authors and Affiliations

  1. NERC Centre for Ecology and Hydrology, Monks Wood, Abbots Ripton, Huntingdon, Cambridgeshire, PE28 2LS, UK

    Roger L. H. Dennis

  2. Institute for Environment, Sustainability and Regeneration, Mellor Building, Staffordshire University, College Road, Stoke on Trent, ST4 2DE, UK

    Roger L. H. Dennis

  3. Istituto Comprensivo Materna Elementere Media Convenevole da Prato, Via 1° Maggio 40, 59100, Prato, Italy

    Leonardo Dapporto

  4. School of Life Sciences, Oxford Brookes University, Headington, Oxford, OX3 0BP, UK

    Tim G. Shreeve

  5. Davies Cottage, Penllyn, Cowbridge, Vale of Glamorgan, CF71 7RQ, UK

    Eddie John

  6. 4 Glykonos Street, Athens, 10675, Greece

    John G. Coutsis

  7. Naturmuseum Südtirol, Bindergasse 1, 39100, Bozen, Italy

    Otakar Kudrna

  8. South Karelia Allergy and Environment Institute, 55330, Tiuruniemi, Finland

    Kimmo Saarinen

  9. Department of Natural Sciences, University of Gävle, 801 76, Gavle, Sweden

    Nils Ryrholm

  10. Applied Psychology, Durham University, Queens Campus, Stockton on Tees, TS17 6BH, UK

    W. R. (Bob) Williams

Authors
  1. Roger L. H. Dennis
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  2. Leonardo Dapporto
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  3. Tim G. Shreeve
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  4. Eddie John
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Correspondence to Roger L. H. Dennis.

Appendices

[n] = number of five indicator species (Pieris brassicae, P. rapae, Colias crocea, Vanessa atalanta; V. cardui) present on island. Islands surveyed at least 15 times

Appendix 1: List of European islands with their locations (longitude and latitude) used in the survey, organised by island group

[n] = number of five indicator species (Pieris brassicae, P. rapae, Colias crocea, Vanessa atalanta; V. cardui) present on island. Islands surveyed at least 15 times

Group 1. Aegean and east Mediterranean islands

East Mediterranean

  • Cyprus (Greece, Turkey) [5] 33.00E 35.00N

  • (Makris 2003; John 2007; John et al. 2006)

Aegean and Aegean arc (Greece and Turkey)

  • Amorgós [0] 25.54E 36.49N

  • Anáfi [0] 25.44E 36.23N

  • Andíparos [1] 25.02E 37.00N

  • Ándros [5] 24.54E 37.49N

  • Astipálea [3] 26.22E 36.32N

  • Áyios Efstrátios [2] 24.58E 39.34N

  • Évvia [5] 23.15E 38.30N

  • Éyina (Aegina) [5] 23.26E 37.45N

  • Folégandros [0] 24.53E 36.37N

  • Foúrni [3] 26.29E 37.36N

  • Gökçeada [3] 25.54E 40.12N

  • Híos (Chios) [5] 26.07E 38.23N

  • Ídhra [3] 23.28E 37.19N

  • Ikaría (Nikaria) [5] 25.40E 37.35N

  • Inoússa [3] 26.14E 38.33N

  • Íos [1] 25.16E 36.44N

  • Kálimnos [5] 26.59E 36.57N

  • Kárpathos (Scarpanto) [5] 27.12E 35.30N

  • Kássos [3] 26.58E 35.22N

  • Kéa [3] 24.20E 37.38N

  • Kíthira [5] 22.59E 36.10N

  • Kíthnos [0] 24.25E 37.25N

  • Kós (Cos) [5] 27.19E 36.53N

  • Kríti (Crete) [5] 25.00E 35.00N

  • Léros [5] 26.50E 37.10N

  • Lésvos (Lesbos) [5] 26.16E 39.16N

  • Límnos (Lemnos) [5] 25.19E 39.53N

  • Megísti (Kastellórizon) [4] 29.35E 36.08N

  • Míkonos [0] 25.22E 37.26N

  • Mílos [5] 24.25E 36.44N

  • Náxos [4] 25.24E 37.06N

  • Níssiros [5] 27.13E 36.35N

  • Páros [5] 25.06E 37.04N

  • Pátmos [5] 26.35E 37.20N

  • Políegos [0] 24.37E 36.46N

  • Psará [1] 27.09E 36.57N

  • Psérimos [1] 27.09E 36.57N

  • Ródhos (Rhodes) [5] 28.00E 36.15N

  • Sámos [5] 26.50E 37.41N

  • Samothráki [5] 25.32E 40.27N

  • Sífnos [5] 24.43E 36.59N

  • Síkinos [0] 25.05E 36.40N

  • Sími [5] 27.51E 36.36N

  • Síros [5] 24.55E 37.26N

  • Skiáthos [5] 23.30E 39.10N

  • Skíros [5] 24.34E 38.55N

  • Skópelos [0] 23.43E 39.07N

  • Spétse (Spetses) [5] 23.09E 37.16N

  • Thássos [5] 24.34E 40.39N

  • Thíra (Santorini) [5] 25.27E 36.24N

  • Tílos [5] 27.18E 36.49N

  • Tínos [5] 25.08E 37.33N

(Gaskin and Littler 1988; Dennis et al. 2000, 2001a, b; Coutsis pers. obs.; Gascoigne-Pees pers. comm.; http://www.butterfly-guide.co.uk/regions/greece/; http://thasos.users.btopenworld.com/butterflies.htm; http://www.chiosnet.gr/tourism/nature/chios_butterflies_moths.htm).

Group 2. Adriatic islands

Adriatic Sea (Croatia)

  • Brac [5] 16.30E 43.20N

  • Cres-Lošinj [5] 14.25E 44.50N

  • Dugi otok [5] 15.02E 44.00N

  • Hvar [5] 16.28E 43.11N

  • Korčula [5] 17.05E 42.56N

  • Kornat [0] 15.20E 43.50N

  • Krk [5] 14.34E 45.02N

  • Mljet [5] 17.30E 42.45N

  • Pag [5] 15.00E 44.50N

  • Rab [5] 14.50E 44.50N

  • Rava [2] 15.05E 44.02N

  • Šolta [2] 16.15E 43.20N

  • Sušak [2] 16.30E 42.45N

  • Unije [5] 14.15E 44.40N

  • Vis [5] 16.11E 43.03N

(Jakšić 1988; Withrington and Verovnik 2008)

Ionian Sea (Greece)

  • Itháki [0] 20.40E 38.25N

  • Kefallinia (Cephalonia) [5] 20.29E 38.13N

  • Kérkira (Corfu) [5] 19.55E 39.38N

  • Levkás [3] 20.30E 38.40N

  • Zákinthos (Zante) [5] 20.54E 37.47N

(Rebel 1910, 1932; Willemse 1980, 1981; Gaskin and Littler 1986; Gaskin 1995; Parker 1996, 2007; Whitehead 1998; Hall et al. 2003; Papapavlou and Katsouni 2008; Olivier pers. comm. [Hipparchia volgensis])

Tremiti Islands (Italy) Adriatic

  • Caprara [3] 15.33E 42.08N

  • San Domino [3] 15.30E 42.06N

  • San Nicola [4] 15.31E 42.07N

(Balletto et al. 2005)

Group 3. West Mediterranean islands

North central Mediterranean

  • Corse (Corsica) (France) [5] 9.00E 42.00N

  • Sardegna (Sardinia) (Italy) [5] 9.00E 40.00N

(Rungs 1988; Lafranchis 2001; Kudrna 2002; Shaw 2002; Jutzeler et al. 2003; Gallet 2003; Brusseaux and Nel 2004; Balletto et al. 2005; Leigheb et al. 2005–2006; Guyot pers. comm.)

Sardinian islands

  • Caprera [5] 9.30E 41.11N

  • Isola Asinara [5] 8.14E 41.05N

  • Isola dei Cavoli [3] 0.33E 39.05N

  • Isola di Maddalena [4] 9.25E 41.13N

  • Isola di Mal di Ventre [3] 8.18E 39.59N

  • Isola di S. Pietro/Panarea [4] 8.15E 39.08N

  • Isola di S. Antioco [5] 8.25E 39.04N

  • Isola la Vacca [3] 8.26E 38.56N

  • Isola S. Maria [5] 9.23E 41.17N

  • Isola Molara [2] 9.45E 40.53N

  • Isola Tavolara [4] 9.43E 40.55N

  • Serpentara [4] 9.38E 39.08N

  • Spargi [4] 9.22E 41.15N

Archipelago Toscano: Ligurian/Tyyrhenian sea (Italy)

  • Capraia [4] 9.54E 43.03N

  • Elba [5] 10.20E 42.49N

  • Gorgona [5] 9.54E 43.25N

  • Giglio [4] 10.54E 42.23N

  • I di Giannutri [3] 11.06E 42.15N

  • M. Angentario [5] 11.11E 42.23N

  • Montecristo [5] 10.18E 42.20N

  • Pianosa [5] 10.06E 42.35N

Golfo di Gaeta/Salerno: Tyrrhenian sea (Italy)

  • Capri [5] 14.13E 40.33N

  • Ischia [5] 13.57E 40.44N

  • Nisida [4] 14.09E 40.49N

  • Ponza [4] 12.58E 40.53N

  • Procida-Vivara [5] 14.01E 40.46N

Æolian Islands: Tyrrhenian Sea (Italy)

  • Lipari [4] 14.58E 38.27N

  • Salina [4] 14.52E 38.34N

  • Stromboli [5] 15.13E 38.47N

  • Ustica [4] 13.12E 38.43N

  • Vulcano [4] 14.58E 38.26N

South central Mediterranean (Italy)

  • Favignana [1] 12.22E 37.56N

  • Lampedusa [5] 12.37E 35.31N

  • Linosa [2] 12.52E 35.53N

  • Pantelleria [5] 11.57E 36.50N

  • Sicilia (Sicily) [5] 14.00E 37.30N

(Balletto et al. 2005; Russell et al. 2005; Dapporto and Cini 2007; Dapporto unpublished data; Dapporto and Dennis in prep., Parenzan and Porcelli 2006; Volpe and Palmieri 2006; Dapporto unpublished data)

Maltese Islands

  • Comino [4] 14.21E 36.01N

  • Gozo [5] 14.15E 36.03N

  • Malta [5] 14.30E 35.00N

(Valletta 1971; Sammut pers. comm.)

Balearic Islands (Spain)

  • Cabrera [2] 2.56E 39.08N

  • Formentera [5] 1.30E 38.41N

  • Ibiza [5] 1.26E 38.54N

  • Mallorca (Majorca) [5] 2.39E 39.35N

  • Menorca (Minorca) [5] 4.15E 39.54N

(Kudrna 2002; García-Barros et al. 2004; Carreras et al. 2004, Honey pers. comm.)

Group 4. South oceanic volcanic islands (Macaronesia)

Azores (Portugal) West

  • Corvo [4] 31.08W 39.41N

  • Flores [4] 31.13W 39.30N

Azores (Portugal) Mid

  • Faial [4] 28.42W 38.35N

  • Graciosa [3] 28.03W 39.03N

  • Pico [3] 28.25W 38.28N

  • Sâo Jorge [3] 28.03W 38.40N

  • Terceira [4] 27.13W 38.43N

Azores (Portugal) East

  • Sâo Miguel [4] 25.27W 37.33N

  • Santa Maria [4] 25.07W 36.58N

(Owen and Smith 1993, Meyer 1993; Olivier and Coutsis 1997; Russell 2003; Vieira 2003; Tennent 2005)

Madeiras (Portugal)

  • Madeira [5] 16.53W 32.38N

  • Porto Santo [4] 16.20W 33.04N

Canary islands (Spain)

  • El Hierro [4] 17.55W 27.48N

  • Fuerteventura [4] 14.02W 28.20N

  • Gran Canaria [4] 15.34W 27.56N

  • La Gomera [5] 17.14W 28.07N

  • Lanzarote [4] 13.38W 29.01N

  • La Palma [5] 17.46W 28.41N

  • Tenerife [5] 16.39W 28.17N

(Owen and Smith 1993, Wiemers 1995; Baldwin 1995; Brunton and Hurst 1998; Meyer 1993; Foster 2000; Wakeham-Dawson et al. 2001, 2002; Gascoigne-Pees et al. 2002; Wakeham-Dawson and Franquinho Aguiar 2003, Fernandez 2004; Tennent 2005)

Group 5. West Atlantic coastal islands

French islands/Bay of Biscay

  • Belle Ile [5] 3.10W 47.19N

  • Ile de Noirmoutier [5] 2.15W 47.01N

  • Ile d’Oléron [5] 1.14W 45.59N

  • Ile de Ré [5] 1.28W 46.09N

  • Ile d’Yeu [5] 2.21W 46.44N

(Coombes pers. comm.; Luquet pers. comm.; Perrein and Guilloton pers. comm.; Lévesque 1994)

Channel Islands (UK)

  • Alderney [5] 2.12W 49.43N

  • Herm [5] 2.27W 49.28N

  • Guernsey [5] 2.35W 49.27N

  • Jersey [5] 2.07W 49.13N

  • Jethou [2] 2.26W 49.27N

  • Sark [5] 2.22W 49.26N

(Dennis and Shreeve 1996; Hull pers. comm.)

Wadden Sea (Netherlands)

  • Ameland [5] 5.46E 53.27N

  • Gried [4] 5.15E 53.15N

  • Rottumeroog and Rottumerplaat [4] 6.35E 53.55N

  • Schiermonnikoog [5] 6.10E 53.28N

  • Terschelling [5] 5.21E 53.24N

  • Texel [5] 4.49E 53.04N

  • Vlieland [5] 40.55E 53.15N

(Tax 1989; Hoentjen and Meijering 1994; Zumkehr 1994)

North Sea (Germany)

  • Amrum [5] 8.20E 54.38N

  • Föhr [1] 8.30E 54.43N

  • Gröde-Appelland [4] 8.38E 54.38N

  • Heligoland [2] 7.53E 54.10N

  • Pellworm [0 ] 8.39E 54.31N

  • Sylt [5] 8.19E 54.53N

(Kolligs 2003, http://www.oag-helgoland.de/beob/aktuell.html).

Off Jutland (Denmark)

  • Fanø [4] 8.20E 55.20N

  • Mandø [4] 8.30E 55.15N

  • Romø [4] 8.30E 55.05N

(Stoltze 1996)

Group 6. British (English, Welsh and Scottish) islands

British islands

  • British mainland [5] 2.20W 54.28N

England

  • Brownsea [5] 1.59W 50.40N

  • Brownsman [4] 1.36W 55.38N

  • Bryher [5] 6.21W 49.57N

  • Great Arthur [0] 6.26W 49.94 N

  • Great Ganilly [3] 6.16W 49.56N

  • Great Innisvouls [0] 6.14E 49.57N

  • Gugh [2] 6.20W 49.54N

  • Hayling [5] 0.59W 50.48N

  • Hilbre [5] 3.10W 53.22N

  • Inner Farne [4] 1.36W 55.38N

  • Lindisfarne [5] 1.48W 55.41N

  • Looe or St George’s [3] 4.27W 50.20N

  • Longstone [1] 1.36W 55.38N

  • Lundy [5] 4.40W 51.11N

  • Menawethan [0] 6.14W 49.56N

  • North Hares [1] 1.36W 55.38N

  • North Wamses [1] 1.37W 55.38N

  • Round [4] 6.19W 49.58N

  • Samson [3] 6.21W 49.55N

  • Sheppey [5] 0.28W 51.25N

  • South Wamses [0] 1.37W 55.37N

  • St Agnes [5] 6.21W 49.54N

  • Staple [3] 1.37W 55.37N

  • St Helens [1] 6.19W 49.58N

  • St Martin’s [5] 6.17W 49.58N

  • St Mary’s Isles of Scilly [5] 6.18W 49.55N

  • St Michael’s Mount [4] 5.29W 50.06N

  • Steepholm [5] 3.07W 51.21N

  • Tean [3] 6.19W 49.58N

  • Thorney island [1] 0.55W 50.49N

  • Tresco [5] 6.20W 49.57N

  • West Wideopens [1] 1.38W 55.36N

  • White Island [0] 6.18W 49.59N

  • Wight [5] 1.18W 52.42N

Wales

  • Anglesey [5] 4.18W 53.16N

  • Bardsey [5] 4.48W 52.46N

  • Caldey [4] 4.42W 51.38N

  • Flatholm [4] 3.07W 51.22N

  • Grassholm [4] 5.29W 51.43N

  • Cardigan Island [0] 4.42W 52.09N

  • Holy Island, off Anglesey [5] 4.39W 53.16N

  • Llanddwyn [5] 4.24W 53.08N

  • Penrhyn-Gwyr [4] 4.20W 51.34N

  • Puffin [5] 4.01W 53.19N

  • Ramsey [4] 5.20W 51.53N

  • Skokholm [5] 5.16W 51.42N

  • Skomer [5] 5.18W 51.45N

  • Walney [5] 3.15W 54.05N

Man

  • Calf of Man [5] 4.46W 54.02N

  • Chicken Rock [1] 4.48W 54.03N

  • Man [5] 4.34W 54.11N

Scotland

  • Ailsa Craig [4] 5.07W 55.16N

  • Arran [5] 5.15E 55.30N

  • Baleshare [2] 7.23W 57.32N

  • Barra [5] 7.30W 56.59N

  • Bass Rock [4] 2.38W 56.05N

  • Benbecula [4] 7.15W 57.26N

  • Berneray nf98 [4] 7.11W 57.43N

  • Berneray nl57 [1] 7.38W 56.47N

  • Bressay [3] 1.05W 60.09N

  • Burnt Islands [2] 5.10W 55.55N

  • Burray [2] 2.54W 58.51N

  • Bute [5] 5.03W 55.51N

  • Calavay [0] 7.15W 57.30N

  • Canna [5] 6.30W 57.03N

  • Cara [3] 5.45W 55.38N

  • Carn nan sgeir [1] 5.22W 57.58N

  • Coll [5] 6.31W 56.38N

  • Colonsay [4] 6.10W 56.05N

  • Crowlin Islands [0] 5.50W 57.20N

  • Cramond [5] 3.20W 55.59N

  • Davaar [3] 5.32W 55.25N

  • Easdale [4] 5.38W 56.18N

  • Eigg [5] 6.10W 56.55N

  • Eilean Furadh Mor [0] 5.42W 57.52N

  • Eilean nan Each [0] 6.15W 56.50N

  • Ensay [2] 7.05W 57.46N

  • Eriskay [4] 7.15W 57.05N

  • Fair Isle [5] 1.38W 59.32N

  • Fetlar[3] 0.52W 60.36N

  • Fiaray [0] 7.26W 57.04N

  • Fidra [3] 2.48W 56.40N

  • Fladday [1] 7.50W 58.10N

  • Flodday [0] 7.21W 56.60N

  • Foula [4] 2.05W 60.08N

  • Fuday [0] 7.23W 57.03N

  • Garvellachs [0] 5.46W 56.15N

  • Gigha [4] 5.45W 55.41

  • Gighay [0] 7.20W 57.01N

  • Glas Eilean [2] 6.38W 57.51N

  • Glunimore [1] 5.33W 55.17N

  • Graemsay [0] 3.17W 58.56N

  • Great Bernera [1] 6.51W 58.13N

  • Great Cumbrae [4] 4.55W 55.46N

  • Grimsay [2] 7.16W 57.30N

  • Gunna [4] 6.44W 56.34N

  • Handa [4] 5.11W 58.23N

  • Harris-Lewis [5] 6.36W 58.05N

  • Heisker [1] 7.40W 57.31N

  • Hellisay [0] 7.21W 57.01N

  • Holy Island, off Arran [3] 5.15W 55.32N

  • Horse [3] 5.20W 57.59N

  • Hoy [4] 3.12W 58.49N

  • Inchcolm [4] 3.18W 56.02N

  • Inchkeith [3] 3.08W 56.02N

  • Inchmarnock [3] 5.09W 55.47N

  • Inchmickery [2] 3.27W 56.01N

  • Iona [4] 6.25W 56.19N

  • Islay [5] 6.17W 55.45N

  • Jura [4] 5.50W 55.57N

  • Kerrera [4] 5.33W 56.24N

  • Killegray [1] 7.05W 57.45N

  • Lady Isle nr Arran [4] 4.44W 55.31N

  • Lismore [1] 5.30W 56.30N

  • Little Bernera [0] 6.52W 58.16N

  • Little Cumbrae [4] 4.57W 55.43N

  • Longa [0] 5.48W 57.44N

  • Longay [2] 5.53W 57.19N

  • Luing [0] 5.38W 56.14N

  • Lunga [1] 5.42W 56.12N

  • May [5] 2.33W 56.12N

  • Minard [2] 5.15W 56.06N

  • Mingulay [0] 7.38W 56.49N

  • Monach Islands [0] 7.38W 57.31N

  • Mousa [1] 1.10W 60.00N

  • Muck [2] 6.15W 56.50N

  • Muldoanich [0] 7.26W 56.55N

  • Mull [4] 6.05W 56.37N

  • Noss [3] 1.05W 60.09N

  • North Rona [1] 5.82W 59.12N

  • North Ronaldsay [4] 2.23W 59.23N

  • North Uist [4] 7.16W 57.37N

  • Orkney mainland [5] 2.58W 58.29N

  • Oronsay [1] 5.56W 56.40N

  • Outer Skerries [2] 0.46W 60.25N

  • Pabay [1] 5.52W 57.16N

  • Pabbay nf88 [0] 7.13W 57.46N

  • Pabbay nl68 [0] 7.34W 56.51N

  • Papa Stour [3] 1.23W 60.20N

  • Pladda [3] 5.06W 55.26N

  • Preist [0] 5.30W 57.58N

  • Raasay [4] 6.03W 57.24N

  • Ronay [0] 7.11W 57.29N

  • Rousay [2] 3.02W 59.10N

  • Rhum [4] 6.17W 57.01N

  • Sanda [3] 5.35W 55.17N

  • Sanday ng20 [3] 6.29W 57.02N

  • Sanday hy63 [3] 2.34W 59.15N

  • Sandray [1] 7.52W 56.89N

  • Scalpay [4] 5.57W 57.18N

  • Scarba [0] 5.42W 56.10N

  • Scarp [1] 7.08W 58.01N

  • Scotasay [0] 6.44W 57.52N

  • Seil [4] 5.37W 56.17N

  • Sgat Mor [1] 5.18W 55.50N

  • Shapinsay [2] 2.34W 59.02N

  • Shiant islands [0] 6.21W 57.54N

  • Sheep [1] 5.57W 55.29N

  • Shetland mainland [5] 1.09W 60.09N

  • Shillay [0] 7.15W 57.48N

  • Skye [4] 6.12W 57.24N

  • Soa [0] 6.76W 56.51N

  • Soay [1] 6.13W 57.08N

  • South Rona [3] 5.59W 57.33N

  • South Ronaldsay [4] 2.57W 58.44N

  • South Uist [4] 7.24W 57.17N

  • Staffa [0] 6.21W 56.26N

  • St Kilda [3] 8.35W 57.49N

  • Sule Skerry [2] 4.24W 59.05N

  • Summer Islands [0] 5.26W 58.00N

  • Tanera Beg [1] 5.26W 58.00N

  • Tanera Mor [1] 5.24W 58.00N

  • Taransay [0] 7.02W 57.54N

  • Tiree [4] 6.49W 56.30N

  • Treshnish Isles [1] 6.26W 56.29N

  • Uinessan [0] 7.29W 56.55N

  • Ulva [2] 6.13W 56.29N

  • Unst [3] 0.52W 60.46N

  • Uyea [1] 0.59W 60.40N

  • Vatersay [1] 7.32W 56.56N

  • West Burra [1] 1.20W 60.05N

  • Westray [2] 2.57W 59.18N

  • Whalsay [3] 0.59W 60.21N

  • Wiay ng23 [0] 6.30W 57.20N

  • Wiay nf84 [0] 7.12W 57.24N

  • Yell [4] 1.05W 60.36N

(Young and Harper 1986; Dean 1990; Dennis and Shreeve 1996, 1997, Pennington 1997; Hulme 1998; Spalding and Tremewan 1998; Dennis et al. 1998a, b; Masters 2000; Jeffcoate 2008; Bowles and Fox 2007; Archer pers. comm.; Beavis pers. comm.; Dennis pers. obs.; Gander pers. comm.; Greatorex-Davies pers. comm.; Jeffcoate pers. comm.; Sandison pers. comm.; Shreeve pers. obs.).

Group 7. Irish islands

  • Ireland [5] 7.56W 53.25N

  • Achill [4] 9.54W 53.56N

  • Aran [3] 9.42W 53.07N

  • Bartragh Island, Killala Bay [0] 9.09W 54.12N

  • Bear [0] 9.54W 51.38N

  • Clare [4]10.00W 53.49N

  • Clear [5] 6.30W 51.26N

  • Cruit near Gola [1] 8.25W 55.01N

  • Dursey [5] 10.12W 51.36N

  • Garinish [3] 9.32W 51.44N

  • Gola [0] 8.20W 55.04N

  • Gorumna [2] 9.40W 53.15N

  • Great Blasket [4] 10.32W 52.05N

  • Great Copeland Belfast [2] 5.31W 54.40N

  • Great Saltee [5] 6.36W 52.00N

  • Great Skellig [1] 10.32W 51.46N

  • Inishbofin [3] (Mayo) 10.12W 53.37N

  • Inishirrer, Gola [0] 8.20W 55.07N

  • Inishmaan [5] 9.35W 53.04N

  • Inishmeane, Gola [0] 8.20W 55.06N

  • Inishmore [5] 9.45W 53.07N

  • Inishshark [0] 10.17W 53.37N

  • Inishsheer [4] 9.31W 53.03N

  • Inishtrahull [4] 7.14W 55.27N

  • Inishturk [0] 10.06W 53.42N

  • Inishvickillane [2] 10.36W 52.20N

  • John’s Copeland [5] 5.32W 54.41N

  • Lambay [4] 6.01W 53.29N

  • Lesser Saltee [4] 6.36W 52.07N

  • Mutton [1] 9.31W 52.48N

  • Omey [1] 10.09W 53.32N

  • Owey Island Donegal [0] 8.26W 55.03N

  • Puffin [2] 10.24W 51.50N

  • Rathlin [5] 6.15W 55.17N

  • Scarrif [2] 10.14W 51.43N

  • Sherkin [5] 9.25W 51.28N

  • Tory [5] 8.13W 55.15N

  • Umfin, Gola [0] 8.21W 55.06N

  • Valencia [3] 10.21W 51.54N

  • Whiddy [3] 9.30W 51.41N

(Lavery 1993; Dennis and Shreeve 1996; Wedd 2004; Rippey pers. comm.; Thymelicus lineola in 2005/2006: http://www.butterflyireland.com/phenology_2006-2005htm#BR).

Group 8. Kattegat islands

Kattegat (Denmark)

  • Agersø [3] 11.15E 55.15N

  • Anholt [4] 11.34E 56.43N

  • Arø [2] 9.40E 55.15N

  • Askø [0] 11.30E 54.52N

  • Avernakø [2] 10.17E 55.02N

  • Bagø [2] 9.45E 55.20N

  • Bjornø [2] 10.15E 55.06N

  • Bogø [5] 12.05E 54.55N

  • Bornholm [5] 14.56E 55.07N

  • Christianso (Ertholmene) [4] 15.12E 55.19N

  • Drejø [3] 10.25E 54.55N

  • Endelave [4] 10.20E 55.40N

  • Falster [5] 12.00E 54.49N

  • Fejø [4] 11.27E 54.50N

  • Femø [4] 11.35E 54.55N

  • Fyn [5]10.20E 55.18N

  • Hesselo [4]11.45E 56.11N

  • Hirscholmene [1] 10.40E 57.30N

  • Hjelm [3] 10.50E 56.10N

  • Laeso [4] 11.01E 57.15N

  • Langeland [5] 10.46E 54.56N

  • Lolland [5] 11.30E 54.57N

  • Lyø [4] 10.10E 55.05N

  • Mon [5] 12.18E 54.59N

  • Nekselø [4] 11.20E 55.45N

  • Nyord [4] 12.10E 55.05N

  • Omø [3] 11.12E 55.10N

  • Ragø [4] 11.20E 54.55N

  • Romsø [3] 10.50E 55.30N

  • Saltholm [4] 12.45E 55.40N

  • Samso [4] 10.36E 55.50N

  • Sejerø [2] 11.10E 55.50N

  • Sjaelland [5] 11.48E 55.28N

  • Skalø [0] 11.25E 54.55N

  • Sprogø [4] 10.55E 55.25N

  • Strynø [1] 10.40E 54.50N

  • Tasinge [4] 10.40E 55.00N

  • Thurø [3] 10.45E 55.05N

  • Tunø [4] 10.30E 55.55N

  • Vejrø [1] 11.25E 55.05N

  • Vejrø1 [1] 10.45E 55.55N

  • Æbelo [2] 10.10E 55.35N

  • Æro [4] 10.20E 54.52N

(Klefbeck 1951; Stoltze 1996)

Kattegat (Sweden)

  • Bohus Malmön [5] 11.19E 58.20N

  • Burholmen [3] 10.59E 58.50N t

  • Hallands Väderö [4] 12.34E 56.25N

  • Nidingen [5] 11.54E 57.18N

  • Nordkoster [4] 11.00E 58.54N

  • Rörö [4] 11.36E 57.46N

  • Saltö [4] 15.34E 56.09N

  • Sydkoster [4] 11.01E 58.52N

  • Tjörn [4] 11.40E 58.05N

  • Ven [5] 12.45E 55.55N

  • Vinga [3] 11.36E 57.37N

(Ryrholm unpublished data)

Group 9. Baltic and Bothnian Sea islands

Gulf of Riga (Estonia)

  • Abruka [4] 22.30E 58.05N

  • Hiiumaa [4] 22.38E 58.40N

  • Kihnu [1] 24.00E 58.05N

  • Muhu [3] 22.40E 58.35N

  • Ruhnu [4] 23.00E 57.48N

  • Saaremaa [5] 22.30E 58.12N

  • Vormsi [3] 22.40E 58.42N

Ålands Islands and Gulf of Finland (Finland)

  • Åland [5] 19.55E 60.15N ✓

  • Brändö main island [3] 21.05E 60.25N ✓

  • Brändö Åva [2] 21.05E 60.26N

  • Brändö Torsholma [0] 21.05E 60.24N

  • Brändö Korsö [1] 21.04E 60.23N

  • Eckerö main island [4] 19.33E 60.13N ✓

  • Eckerö Signilskär [2] 19.36E 60.12N

  • Föglö main island [4] 20.28E 60.00N ✓

  • Föglö Överö/ Ulversö [3] 20.32E 60.05N ✓

  • Föglö Finnholma [2] 20.32E 60.04N

  • Föglö Jyddö [3] 20.32E 60.04N ✓

  • Föglö Nötö [4] 20.31E 60.04N ✓

  • Föglö Vargskär [1] 20.33E 60.05N

  • Föglö Bänö [0] 20.34E 60.04N

  • Houtskar [4] 21.05E 60.10N (c. 3-4km2) ✓

  • Isosaari [4] 25.00E 60.08N (1km2; 3km isolation) ✓

  • Kumlinge main island [3] 20.45E 60.15N ✓

  • Kumlinge Seglinge [1] 20.43E 60.13N

  • Kökar main island [4] 20.58E 59.56N ✓

  • Kökar Kyrkogårdsö [0] 20.56E 59.55N

  • Sottunga main island [3] 20.40E 60.06N✓

  • Sottunga Husö [3] 20.47E 60.03N✓

  • Sund Prästö [0] 20.18E 60.14N

  • Vårdö main island [3] 20.20E 60.15N✓

(Nordstrøm 1955; Marttila et al. 2001; Laasonen and Laasonen 1991, Bruun 1992; Kesküla 1992; Huldén pers. comm.; Saarinen unpublished data; ✓ used in island-focussed analysis as having been visited > 15 times)

Baltic Sea (Sweden)

  • Alnön [4] 17.30E 62.46N

  • Björn [4] 17.59E 60.38N

  • Bjuröklubb [2] 21.34E 64.28N

  • Blå Jungfrun [4] 16.46E 57.16N

  • Brämön [3] 17.42E 62.12 N

  • Brändön [3] 22.19E 65.41N

  • Eggegrund [4] 17.33E 60.44N

  • Fagerön [4] 18.27E 60.13N

  • Frevisören [3] 23.24E 65.45N

  • Gotland [5] 18.28E 57.32N

  • Gotska Sandon [4] 19.25E 58.25N

  • Gräsö [4] 18.30E 60.22N

  • Haparanda Sandskär [3] 23.44E 65.20N

  • Holmön [3] 20.52E 63. 47N

  • Hornslandet [4] 17.28E 61.40N

  • Iggön [4] 17.18E 60.53N

  • Inre Hamnskär [4] 19.19E 59.44N

  • Landsort [4] 17.52E 58.45N

  • Måssten [4] 18.51E 60.11N

  • Munkö [4] 18.43E 59.14N

  • Norra Finnö [4] 16.51E 58.19N

  • Obbolaön [3] 20.18E 63.42N

  • Öland [5] 16.40E 56.50N

  • Ornö [4] 18.24E 59.03N

  • Örskär [4]18.23E 60.19N

  • Raggarön [4] 18.35E 60.12N

  • Rävsön [3] 18.28E 62.55N

  • Revsudden [5] 16.27E 56.46N

  • Runmarö [4] 18.45E 59.16N

  • Seskarö [3] 23.44E 65.44N

  • Singö [4] 18.45E 60.10N

  • Sladö [4] 15.24E 56.08N

  • Sturkö [5] 15.42E 56.05N

  • Svartklubben [4] 18.49E 60.10N

  • Ulvön [3] 18.38E 63.01N

  • Utklippan [3] 15.47E 56.01N

  • Utlängan [4] 15.50E 56.02N

  • Utö [4] 18.20E 58.56N

(Gustafsson 1987; Ryrholm unpublished data)

Group 10. Norwegian and North Atlantic islands

  • Andørja (Andøy) [0] 17.15E 68.50N

  • Askøy [1] 5.01E 60.25N

  • Austvågøy (Vågøy) [1] 14.30E 68.20N

  • Bjarkøy [0] 16.30E 69.00N

  • Bømlo [0] 5.15E 59.45N

  • Boroy [3] 10.33E 59.52N

  • Bronnoya [1] 10.32E 59.51N

  • Dønna [0] 12.30E 66.10N

  • Finnøy [1] 5.50E 59.10N

  • Frøya [0] 8.30E 63.45N

  • Hadseløya (Hadsel) [0] 14.30E 68.30N

  • Håoya [0] 10.34E 59.41N

  • Hinnø (Lødingen) [1] 15.45E 68.30N

  • Hitra [1] 8.30E 63.30N

  • Holsnøy (Meland) [0] 5.05E 60.35N

  • Huftarøy (Austevoll) [0] 5.15E 60.03N

  • Huser [0] 10.55E 59.05N

  • Karlsøy [0] 20.00E 70.00N

  • Kirkeøy (Hvaler) [3] 11.02E 59.05N

  • Kristiansund [0] 7.45E 63.05N

  • Kvåløya (Hammerfest) [0] 24.00E 70.35N

  • Kvitsøy [1] 5.25E 59.05N

  • Langoya, Våle [2] 10.23E 59.29N

  • Lurøy [0] 12.45E 66.25N

  • Moskenesøya [0] 12.45E 68.00N

  • Øksnes [0] 14.55E 68.50N

  • Ostoya [3] 10.34E 59.52N

  • Rauer [0] 10.41E 59.13N

  • Rennesøy [0 ] 5.45E 59.07N

  • Rølla (Ibestad) [0] 17.00E 68.45N

  • Senja (Tranøy, Berg, Torsken) [0] 17.00E 69.20N

  • Smøla [0] 8.00E 63.02N

  • Sotra-Store (Sund, Fjell, Øygarden) [1] 5.05E 60.17N

  • Stord [0] 5.25E 59.52N

  • Sula (Solund) [0] 6.05E 62.25N

  • Sula [0] 4.55E 61.08N

  • Tjeldøya (Tjeldsund) [0] 16.10 68.30N

  • Tjøme [4] 10.25E 59.10N

  • Tofteholmen [2] 10.33E 59.30N

  • Træna [0] 12.02E 66.28N

  • Tromøy [5] 8.53E 58.28N

  • Tysnesøy [1] 5.35E 60.00N

  • Utsira [0] 4.50E 59.20N

  • Vega [0] 11.55E 65.35N

  • Vestvågøy [1] 13.45E 68.15N

Faeroes (Denmark)

  • Streymoy [5] 6.47W 62.02N

Iceland

  • Iceland [2] 18.00W 65.00N

(Williams 1958; Nagypal pers. comm.; http://www.toyen.uio.no/norlep/; Ryrholm unpublished data; http://www.faroenature.net/UserFiles/File/FFFFFF_1998.pdf (Jensen and Patursson A, pp 9–12)).

Appendix 2: List of butterfly species used in study of European islands*

Owing to uncertainty of the records, various taxa have been amalgamated in the listing of species. For example Aricia artaxerxes (Fabricius, 1793) includes reported sightings of A. allous Geyer, 1837 and Pieris daplidice (Linnaeus, 1758) includes P. edusa (Fabricius, 1777).

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Dennis, R.L.H., Dapporto, L., Shreeve, T.G. et al. Butterflies of European islands: the implications of the geography and ecology of rarity and endemicity for conservation. J Insect Conserv 12, 205–236 (2008). https://doi.org/10.1007/s10841-008-9148-3

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