Abstract
The biology and ecology of the European species of the genus Pyrgus Hübner are revised and novel records on theirfoodplants, parasitoids and immature stages are provided. The DNA barcoding technique was used in some cases for theidentification of eggs or larvae, showing the effective use of this tool for a species level characterization within the genus.We thoroughly describe morphological details from the eggs, last instar larvae and pupae using scanning electronmicroscope images. With the studied morphological and ecological characters we performed a cladistic analysis whoseresults are highly coincident with classifications based on adult morphology. Therefore, the characters of immature stagesprove to be valuable for the taxonomical study of the genus. Our results point out the possibility of combining the studiedcharacters, not only among them, but also with other datasets (adult morphology, molecular characters) in order to infer phylogenetic relationships within the genus Pyrgus.References
Abadjiev, S. (2001) An Atlas of the Distribution of the Butterflies in Bulgaria (Lepidoptera: Hesperioidea & Papilionoidea). Pensoft Publishers, Sofia, 335 pp.
Anthos (2011) Sistema de información de las plantas de España. Real Jardín Botánico, CSIC Fundación Biodiversidad, Madrid. Avalaible from: http://www.anthos.es/ (accessed September 2011).
Aussem, B. (1980) Die Präimaginalstadien der Gattung Pseudochazara De Lesse, 1951. Teil 1: Pseudochazara graeca (Staudinger, 1870). Mitteilungen Münchener Entomologischen Gesellschaft, 69, 1–7.
Bink, F.A. (1992) Ecologische Atlas van de Dagvlinders van Noordwest-Europa. 2. Auflage. Schuyt & Co, Haarlem, 512 pp.
Bink, F.A. & Weidemann, H.J. (1995) Familie Hesperiidae, Dickkopffalter. In: Weidemann, H.J. (Ed.), Tagfalter: beobachten, bestimmen. 2. Auflage. Naturbuch-Verlag, Augsburg, pp. 586–631.
Brockmann, E., Thust, R. & Kristal, P.M. (1996) Zur Biologie von Pyrgus warrenensis (Verity 1928) (Lepidoptera: Hesperiidae). Nachrichten des Entomologischen Vereins Apollo, 17 (2), 183–189.
BUTT (Butterflies Under Threat Team) (1986) The management of chalk grassland for butterflies. Focus on Nature Conservation, 17, 1–80.
Cates, R.G. (1980) Feeding patterns of monophagus, oligophagus and polyphagus insect herbivores: the effect of resource abundance and plant chemestry. Oecologia, 46, 22–31.
Christensen, P.R. (2000) The effects of grazing on the butterfly fauna in Denmark. PhD thesis dissertation, Department of Zoology, University of Aarhus, Aarhus, 112 pp.
Cottrell, C.B. (1984) Aphytophagy in butterflies: its relationship to myrmecophily. Zoological Journal of the Linnean Society, 80, 1–57.
Courtney, S.P. (1984) Habitat versus foodplant selection. In: Vane-Wright, R.I., Ackery, P.R. (Eds.), The biology of butterflies. Academic Press, London, pp. 89–90.
De Jong, R. (1972) Systematics and geographic history of the genus Pyrgus in the palaearctic region. Tijdschrift voor Entomologie, 115, 1–121.
De Jong, R. (1987) Superspecies Pyrgus malvae (Lepidoptera: Hesperiidae) in the East Mediterranean, with notes on phylogenetic and biological relationships. Zoologische Mededelingen, 61 (34), 483–500.
Drummond, A.J., Ashton, B., Buxton, S., Cheung, M., Cooper, A., Duran, C., Field, M., Heled, J., Kearse, M., Markowitz, S., Moir, R., Stones-Havas, S., Sturrock, S., Thierer, T., Wilson, A. (2011) Geneious v5.4. Biomatters Ltd, Auckland. Available from: http://www.geneious.com/
Ebert, G. & Rennwald, E. (1991) Die Schmetterlinge Baden-Württembergs 2: Tagfalter II. Verlag Eugen Ulmer, Stuttgart, 535 pp.
Efron, B. (1979) Bootstrap methods: Another look at the jack-knife. Annals of Statistics, 7, 1–26.
Evans, W.H. (1953) A Catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum (Natural History). Part III. Pyrginae. Section II. British Museum, London, 246 pp. + pls. 26–53.
Ferreira, T.A. & Rasband, W. (2010) The ImageJ User Guide, Version 1.43. Avalaible from http://rsbweb.nih.gov/ij/docs/user-guide.pdf/
Fiedler, K. (1988) The preimaginal organs of Lycaena tityrus (Poda, 1761) and Polyommatus coridon (Poda, 1761) (Lepidoptera: Lycaenidae)–a comparison. Nota Lepidopterologica, 11 (2), 100–116.
Fiedler, K. (1991) Systematic, evolutionary, and ecological implications of myrmecophily within the Lycaenidae (Insecta: Lepidoptera. Papilionoidea). Bonner Zoologische Monographien, 31, 1–210.
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial Cytochrome C oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
Forster, W. & Wohlfahrt, T.A. (1955) Die Schemetterlinge Mitteleuropas, Bd. II, Tagfalter. Franckh, Stuttgart, 126 pp.
García-Barros, E. & Martín, J. (1991) Immature stages of Hipparchia Fabricius and the systematics of the “Satyrus series” (Lepidoptera: Nymphalidae: Satyrinae). Systematic Entomology, 16, 407–426.
García-Barros, E. & Martín, J. (1994) Morfología de los tubérculos mesotorácicos de las pupas de algunos lepidopteros (Lepidoptera: Nymphalidae, Hesperiidae, Sphingidae). SHILAP Revista de Lepidopterología, 22 (87), 249–258.
García-Barros, E., Munguira, M.L., Martín, J., Romo, H., García-Pereira, P. & Maravalhas, E.S. (2004) Atlas de las Mariposas Diurnas de la Península Ibérica y Baleares (Lepidoptera: Papilionoidea & Hesperioidea). Monografías SEA (11), Zaragoza, 228 pp.
Goloboff, P. (1999) NONA (NO NAME) Ver. 2. Published by the author, Tucumán (computersoftware).
Gómez-Bustillo, M.R. & Fernández-Rubio, F. (1974) Mariposas de la Península Ibérica. Ropalóceros II. ICONA, Madrid, 258 pp.
Gros, P. (1998) Eiablage und Futterpflanzen der Falter der Gattung Pyrgus Hübner, 1819 im Bundesland Salzburg (Lepidoptera: Hesperiidae, Pyrginae). Zeitschrift der Arbeitsgemeinschaft österreichischer Entomologen, 50, 29–36.
Guillaumin, M. (1964) Les espéces françaises de genre Pyrgus Hübner avec clé détermination d’apres les genitalia des mâles (Lep. Hesperiidae). Alexanor, 3, 293–305.
Guillaumin, M. & Descimon, H. (1976) La notion d'espèce chez les Lépidoptères. In: Bocquet, C., Génermont, J. &. Lamotte, M. (Eds.), Les problèmes de l'espèce dans le règne animal. Société zoologique de France, Paris, pp. 129–201.
Hebert, P.D.N., Cywinska, A., Ball, S.L., deWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society (London), Series B (Biological Sciences), 270, 313–321.
Henriksen, H.J. & Kreutzer, I. (1982) The Butterflies of Scandinavia in Nature. Apollo Books, Stenstrup, 455 pp.
Hernández-Roldán, J.L., Munguira, M.L. & Martín, J. (2009) Ecology of a relict population of the vulnerable butterfly Pyrgus sidae on the Iberian Peninsula (Lepidoptera: Hesperiidae). European Journal of Entomology, 106 (4), 611–618.
Hernández-Roldán, J.L., Murria, C., Romo, H., Talavera, G., Zakharov, E., Hebert, P.D.N & Vila, R. (2011a) Tracing the origin of disjunct distributions: a case of biogeographical convergence in Pyrgus butterflies (Insecta: Lepidoptera: Hesperiidae). Journal of Biogeography, 38, 2006–2020.
Hernández-Roldán, J.L., Vicente, J.C. & Munguira, M.L. (2011b) Historia natural de Pyrgus cacaliae (Rambur, 1839) en la Península Ibérica (Lepidoptera: Hesperiidae). SHILAP Revista de Lepidopterología, 153, 61–73.
Higgins, L.G. & Riley, N.D. (1970) A field guide to the butterflies of Britain and Europe. Collins, London, 380 pp.
Kauffmann, G. (1954) Beobachtungen über die ersten Stände einiger alpiner Pyrginae. Zeitschrift der Wiener Entomologischen Gesellschaft, 39, 23–28.
Kudrna, O., Harpke, A., Lux, K., Pennerstorfer, J., Schweiger, O., Settele, J. & Wiemers, M. (2011) Distribution atlas of butterflies in Europe. Gesellschaft für Schmetterlingschutz, Halle, 576 pp.
Lafranchis, T. (2000) Les papillons de jour de France, Belgique et Luxembourg et leurs chenilles. Biotope, Meze, 448 pp.
LSPN (1999) Les papillons et leurs biotopes. Tome 2. Pronatura–Ligue suisse pour la protection de la nature, Bâle, 667 pp.
Malicky, H. (1969) Versuch einer Analyse der ökologischen Beziehungen zwischen Lycaeniden (Lepidoptera) und Formiciden (Hymenoptera). Tijdschrift voor Entomologie, 112, 213–298.
Manley, W.B.L. & Allcard, H.G. (1970) A field guide to the butterflies and burnets of Spain. E.W. Classey, Hampton, 192 pp.
Munguira, M.L. (1989) Biología y biogeografía de los licénidos ibéricos en peligro de extinción. PhD thesis dissertation, Servicio de Publicaciones, Universidad Autónoma de Madrid, Madrid, 462 pp.
Munguira, M.L., García-Barros, E. & Martín, J. (2009) Butterfly herbivory and larval ecology. In: Settele, J., Shreeve, T., Konvicka, M. & Van Dyck, H. (Eds.), Ecology of Butterflies in Europe. Cambridge University Press, Cambridge, pp. 43–54.
Nel, J. (1984) Note sur Pyrgus sidae Esper: sa plante-hôte et son cycle biologique en Provence (Lep. Hesperiidae). Alexanor, 13 (6), 275–281.
Nel, J. (1985a) Note sur l’écologie et la biologie de Pyrgus bellieri Obth. en Provence et dans le Briançonnais. Comparaison avec P. alveus Hb. Alexanor, 14 (1), 3–7.
Nel, J. (1985b) Note sur la répartition, les plantes-hôtes et le cycle de développement des Pyrginae en Provence (Lep. Hesperiidae). Alexanor, 14 (2), 51–63.
Öckinger, E. (2006) Possible metapopulation structure of the threatened butterfly Pyrgus armoricanus in Sweden. Journal of Insect Conservation, 10, 43–51.
Pamperis, L.N. (2009) The Butterflies of Greece. Editions Pamperis, Athens, 768 pp.
Patočka, J. & Turčáni, M. (2005a) Lepidoptera pupae. Central European Species. Text volume. Apollo books, Stenstrup, 542 pp.
Patočka, J. & Turčáni, M. (2005b) Lepidoptera pupae. Central European Species. Plate volume. Apollo books, Stenstrup, 321 pp.
Picard, J. (1950) Lépidoptères Hesperiidae des départaments de l´Ille-et-Vilaine et des Côtes-du-Nord. Entomologiste, 5, 163–169.
Pierce, N. (1983) The ecology and evolution of symbioses between lycaenid butterflies and ants. PhD thesis dissertation, Harvard University, Cambridge, x + 274 pp.
Powell, H. (1911) A contribution to the life history of Hesperia sidae, Esp. Transactions of the Entomological Society of London, 3, 563–576.
Reverdin, J. (1912) Notes sur le genre Hesperia. Bulletin de la Société Lépidoptérologique de Genève, 3, 141–172. + pls. 16–19.
Shaw, M.R., Stefanescu, C. & Van Nouhuys, S. (2009) Parasitoids of European butterflies. In: Settele, J., Shreeve, T., Konvicka, M. & Van Dyck, H. (Eds.), Ecology of Butterflies in Europe. Cambridge University Press, Cambridge, pp. 130–156.
Simon, C., Frati, F., Beckebach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–701.
Stefanescu, C., Peñuelas, J. & Filella, I. (2007) Les papallones com a bioindicadores dels hàbitats a Catalunya: l’exemple dels prats de dall i les pastures del Parc Natural dels Aiguamolls de l’Empordà. Butlletí de la Institució Catalana d’Història Natural, 73 (2005), 139–162.
Székely, L. (2008) The butterflies of Romania / Fluturii de zi din România. Brasov County History Museum, Brastar Print, Brasov, 305 pp.
Thiele, K. (1993) The holy grail of the perfect character: the cladistic treatment of morphometric data. Cladistics, 9, 275–304.
Thust, R., Voigt, M., & Brockmann, E. (1997) Zur rasterelektronenmikroskopischen Morphologie der Eier europäischer Pyrginae (Lepidoptera: Hesperiidae). Nachrichten des Entomologischen Vereins Apollo, 18 (2/3), 109–128.
Tolman, T. & Lewington, R. (1997) Collins Field Guide Butterflies of Britain & Europe. Harper Collins, London, 320 pp.
Tshikolovets, V. (2011) Butterflies of Europe and the Mediterranean Area. Tshikolovets Publications, Pardubice, 544 pp.
Van Swaay, C., Cuttelod, A., Collins, S., Maes, D., López Munguira, M., Sasic, M., Settele, J., Verovnik, R., Verstrael, T., Warren, M., Wiemers, M. & Wynhof, I. (2010) European Red List of Butterfies. Publications Office of the European Union, Luxembourg, 47 pp.
Verity, R. (1940) La Farfalle diurne d´Italia 1. Considerazioni generali. Superfamiglia Hesperides. Ed. Marzocco, Firenze, 34+131pp.
Vicente, J.C. & Hernández-Roldán, J.L. (2007) Guía de las Mariposas Diurnas de Castilla y León. Náyade Editorial, Medina del Campo, 280 pp.
Wagner, W. (2002) Zur Ökologie von Pyrgus trebevicensis (Warren, 1926) und Pyrgus alveus Hübner [1803]) (Lepidoptera: Hesperiidae) auf der Schwäbischen Alb (Baden-Württemberg). Entomologische Zeitschrift, 112, 145–156.
Wagner, W. (2003) Beobachtungen zur Biologie von Pyrgus andromedae (Wallengren, 1853) und P. cacaliae (Rambur, 1840) in den Alpen (Lepidoptera: Hesperiidae). Entomologische Zeitschrift, 113, 346–353.
Wagner, W. (2006) Die Gattung Pyrgus in Mitteleuropa und ihre Ökologie- Larvalhabitate, Nährpflanzen und Entwicklungszyklen. In: Fartmann, T. & Hermann, G. (Eds.): Larvalökologie von Tagfaltern und Widderchen in Mitteleuropa. Abhandlungen aus dem Westfälischen Museum für Naturkunde. Heft 68 (3/4), 83–122.
Wagner, W. (2009a) Zur Ökologie von Pyrgus cinarae (Rambur 1839) nebst Beobachtungen zu Spialia phlomidis (Herrich-Schäffer 1845) - Larvalhabitat, Präimaginalstadien und Entwicklungszyklus (Lepidoptera: Hesperiidae). Nachrichten des Entomologischen Vereins Apollo, 29 (4), 199–204.
Wagner, W. (2009b) Zur Freiland-Raupennahrung von Pyrgus onopordi (Rambur 1839) in Andalusien (Lepidoptera: Hesperiidae). Nachrichten des Entomologischen Vereins Apollo, 30 (1/2), 1–4.
Warren, B.C.S. (1926) Monograph of the tribe Hesperiidi (European species) with revised classification of the subfamily Hesperiinae (Palaearctic species) based on the genital armature of the males. Transactions of the Entomological Society of London, 74, 1–170.
Warren, A.D. (2006) The higher classification of the Hesperiidae (Lepidoptera: Hesperioidea). PhD thesis dissertation, Oregon State University, Oregon, 458 pp.
Warren, A.D., Ogawa, J.R. & Brower, A.V.Z. (2008) Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea). Cladistics, 24, 642–676.
Warren, A.D., Ogawa, J.R. & Brower, A.V.Z. (2009) Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomology, 34, 467–523.
Wiens, J. (2001) Character analysis in morphological phylogenetics: problems and solutions. Systematic Biology, 50, 689–699.
Wiklund, C. (1975) The evolucionary relationship between adult oviposition preferences and larval host plant range in Papilio machaon. Oecologia 18, 185–197.