Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 4402 No. 3: 29 Mar. 2018
Type: Article
Published: 2018-03-29
DOI: 10.11646/zootaxa.4402.3.1
Page range: 401–442
Abstract views: 115
PDF downloaded: 6

Antennal ultrastructure of Leptidea Billberg, 1820 (Pieridae: Dismorphiinae: Leptideini) and its taxonomic implications

Museo de Zoología, Depto. Biología Evolutiva Fac. Ciencias, UNAM, Cd. México, 04510. National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC. 20013–7012, USA.
Museo de Zoología, Depto. Biología Evolutiva Fac. Ciencias, UNAM, Cd. México, 04510. National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC. 20013–7012, USA.
Lepidoptera auriculate basiconic chaetica coeloconic pseudosulci sensilla sulci trichoid trisulcate

Abstract

We present the results of the first examination and comparative analysis of the ultrastructure of the antennae in seven species of Leptidea Billberg, 1820 (Pieridae). The results add further support to the hypothesis that the subfamily Dismorphiinae is composed of the tribes Leptideini and Dismorphiini. We summarize the results in a table and discuss the ultrastructure similarities and differences between the two tribes. Following a previously proposed framework, we trace the progression of character states and discuss the significance of the results in the context of varying phylogenetic hypotheses for Leptidea, with special emphasis on biogeography. We include a brief description of the terms we employ to described the main characters and types of sensilla of the antennal club of Leptidea. Remarkable antennal sexual dimorphism is noted in L. gigantea.

 

References

  1. Amiet, J.L. (2004) Séparation de niches écologiques chez deux espéces jumelles sympatriques de Leptidea (Lepidoptera, Pieridae). Revue d’Ecologie (Terre et vie), 59, 433–455.

    Asmussen, C.B. & Liston, A. (1998) Chloroplast DNA characters, phylogeny, and classification of Lathyrus (Fabaceae). American Journal of Botany, 85 (3), 387–401.
    https://doi.org/10.2307/2446332

    Bolshakov, L. (2004) The new taxa of genus Leptidea Billberg, 1820 (Lepidoptera: Pieridae) from Asians mountains regions. Bulletin of the Moscow Society of Naturalists, 109 (2), 78–83. [in Russian]

    Braby, M.F. & Trueman, J.W.H. (2006) Evolution of larval host plant associations and adaptive radiation in pierid butterflies. Journal of Evolutionary Biology, 19 (5), 1677–1690.
    https://doi.org/10.1111/j.1420-9101.2006.01109.x

    Braby, M.F., Vila, R. & Pierce, N.E. (2006) Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography. Zoological Journal of the Linnean Society, 147 (2), 239–275.
    https://doi.org/10.1111/j.1096-3642.2006.00218.x

    Castro-Gerardino, D.J. & Llorente-Bousquets, J. (2017) Comparative exploration of antennae in Pseudopontia, and antennal clubs of the tribes Leptideini and Dismorphiini (Lepidoptera: Pieridae). Zootaxa, 4347 (3), 401–445.
    https://doi.org/10.11646/zootaxa.4347.3.1

    Cheong, S.W. (1990) Comparative morphology and phylogeny on female internal genitalica of the Pieridae (Lepidoptera). Ph. D. dissertation. Kyungpook National University, Taegu, 9 pp.

    Cheong, S.W., Lee, C.E. & Park, H.C. (1990) A microscopic study on the bursa copulatrix of Korean Pieridae (Lepidoptera). Esakia, Special Issue, 1, 167–172.

    Churkin, S.V. (2004) А new species of Leptidea Billberg, 1820 (Lepidoptera, Pieridae) from Peter the Great Range. Helios, 5, 137–141.

    Clench, H.K. (1955) Revised classification of the butterfly family Lycaenidae and its allies. Annals of the Carnegie Museum, 33, 261–274.

    Cupedo, F. & Hoen, F.W. (2006) Leptidea sinapis and Leptidea reali (Lepidoptera: Pieridae) in The Netherlands. Entomologische Berichten, 66 (4), 118–123.

    Dincă, V., Lukhtanov, V.A., Talavera, G. & Vila, R. (2011) Unexpected layers of cryptic diversity in wood white Leptidea butterflies. Nature Communications, 2 (324), 1–8.
    https://doi.org/10.1038/ncomms1329

    Dincă, V. & Vila, R. (2011) Biodiversitat críptica i espècies falses: papallones que enganyen els científics. Omnis Cellula, 27, 22–27.

    Dincă, V., Wiklund, C., Lukhtanov, V.A., Kodandaramaiah, U., Norén, K., Dapporto, L., Wahlberg, N., Vila, R. & Friberg, M. (2013) Reproductive isolation and patterns of genetic differentiation in a cryptic butterfly species complex. Journal of Evolutionary Biology, 26 (10), 2095–2106.
    https://doi.org/10.1111/jeb.12211

    Ding, C. & Zhang, Y. (2017) Phylogenetic relationships of Pieridae (Lepidoptera: Papilionoidea) in China based on seven gene fragments. Entomological Science, 20 (1), 15–23.
    https://doi.org/10.1111/ens.12214

    Ehrlich, P. (1958) The comparative morphology, phylogeny and higher classification of the butterflies (Lepidoptera: Papilionoidea). University of Kansas Science Bulletin, 39, 305–370.

    Faucheux, M.J. (1999) Biodiversité et unité des organes sensoriels des Insectes Lépidoptères. Bulletin de la Société des Sciences Naturelles de l'Ouest de la France,Supp. Hors Série, 1–294.

    Fracasso A. (2002) Leptidea reali (Reissinger, 1989): Note preliminari sulla distribuzione nel nord-est dell’Italia (Lepidoptera, Pieridae). Bollettino del Museo civico di Storia Naturale di Venezia, 53, 149‒154.

    Freese, A. & Fiedler, K. (2002) Experimental evidence for specific distinctness of the two wood white butterfly taxa, Leptidea sinapis and L. reali (Pieridae). Nota Lepidopterologica, 25 (1), 39–59.

    Friberg, M. (2007) A difference in pupal morphology between the sibling species Leptidea sinapis and L. reali (Pieridae). Nota Lepidopterologica, 30 (1), 61–64.

    Friberg, M., Bergman, J., Kullberg, M., Wahlberg, N. & Wiklund, C. (2008a) Niche separation in space and time between two sympatric sister species —a case of ecological pleiotropy. Evolutionary Ecology, 22 (1), 1–18.
    https://doi.org/10.1007/s10682-007-9155-y

    Friberg, M., Leimar, O. & Wiklund, C. (2013) Heterospecific courtship, minority effects and niche separation between cryptic butterfly species. Journal of Evolutionary Biology, 26 (5), 971–979.
    https://doi.org/10.1111/jeb.12106

    Friberg, M., Olofsson, M., Berger, D., Karlsson, B. & Wiklund, C. (2008b) Habitat choice precedes host plant choice —niche separation in a species pair of a generalist and a specialist butterfly. Oikos, 117 (9), 1337–1344.
    https://doi.org/10.1111/j.0030-1299.2008.16740.x

    Friberg, M., Vongvanich, N., Borg-Karlson, A.K., Kemp, D.J., Merilaita, S. & Wiklund, C. (2007) Female mate choice determines reproductive isolation between sympatric butterflies. Behavioral Ecology and Sociobiology, 62 (6), 873–886.
    https://doi.org/10.1007/s00265-007-0511-2

    Friberg, M. & Wiklund, C. (2007) Generation-dependent female choice: behavioral polyphenism in a bivoltine butterfly. Behavioral Ecology, 18 (4), 758–763.
    https://doi.org/10.1093/beheco/arm037

    Friberg, M. & Wiklund, C. (2009) Host plant preference and performance of the sibling species of butterflies Leptidea sinapis and Leptidea reali: a test of the trade-off hypothesis for food specialization. Oecologia, 159 (1), 127–137.
    https://doi.org/10.1007/s00442-008-1206-8

    Fumi, M. (2008) Distinguishing between Leptidea sinapis and L. reali (Lepidoptera: Pieridae) using a morphometric approach: impact of measurement error on the discriminative characters. Zootaxa, 1819, 40–54.

    Gallo, F., Beretta, S., Salogni, G. & Bonato, L. (2015) Partial separation of climatic niche between two cryptic species of Leptidea (Lepidoptera: Pieridae) in the Alpine-Adriatic region. Italian Journal of Zoology, 82 (4), 573–581.
    https://doi.org/10.1080/11250003.2015.1061614

    Ghiradella, H. (1998) Hairs, bristles, and scales. In: Harrison, F.W. & Locke, M. (Ed.), Microscopic Anatomy of Invertebrates. Vol. 11A. Insecta. Wiley-Liss, New York, pp. 257–287.

    Gorbunov, P.Y. (2001) Butterflies of Russia: classification, genitalia keys for identification (Lepidoptera: Hesperioidea and Papilionoidea). Edt. Thesis, Ekaterinburg, 320 pp.

    Grote, R. (1900) The descent of the pierids. Proceedings of the American Philosophical Society, 39 (161), 4–67.

    Hallberg, E., Hansson, B.S. & Löfstedt, C. (2003) Sensilla and proprioceptors. In: Kristensen, N.P. (Ed.), Handbook of zoology. Vol. IV. Part 36. Lepidoptera 2. Moths and butterflies. Vol. 2. Morphology, physiology and development. Walter de Gruyter, Berlin, pp.267–288. https://doi.org/10.1515/9783110893724.267

    Häuser, C.L. (1993) Die inneren weiblichen Genitalorgane der Tagfalter (Rhopalocera): Vergleichende Morphologie und phylogenetische Interpretation (Insecta, Lepidoptera). [The internal female genital organ in butterflies (Rhopalocera): comparative morphology and phylogenetic intepretation (Insecta, Lepidoptera)]. Zoologische-Jahrbucher, Abeteilung fur-Systematik, Okologie und-Geographie-Tiere, 120 (4), 389–439.

    Higgins, L.G. (1975) The classification of European butterflies. Collins, London, 320 pp.

    Higgins, L.G. & Riley, N.D. (1993) A field guide to the butterflies of Britain and Europe. Harper Collins, London, 380 pp.

    Jeffcoate, S. (2006) Seasonal variation in the use of vegetation resources by Leptidea sinapis (Linnaeus, 1758) (Lepidoptera: Pieridae), a multivoltine species in southern Britain: implications for its conservation at the edge of its range and in the context of climate change. Entomologists Gazette, 57 (2), 69–82.

    Jong de, R. & Van Achterberg, C. (2007) Global disjunctions and flying insects. In: Renema, W. (Ed.), Biogeography, Time, and Place: Distributions, Barriers, and Islands. Springer, Dordrecht, pp. 5–44.
    https://doi.org/10.1007/978-1-4020-6374-9_1

    Jong de, R., Vane-Wright, R.I. & Ackery, P.R. (1996) The higher classification of butterflies (Lepidoptera): problems and prospects. Insect Systematics & Evolution, 27 (1), 65–101.
    https://doi.org/10.1163/187631296x00205

    Jordan, K. (1898) Contributions to the Morphology of Lepidoptera. Novitates Zoologicae, 5, 374–415.

    Keil, T.A. (1997) Comparative morphogenesis of sensilla: a review. International Journal of Insect Morphology and Embryology, 26 (3), 151–160.
    https://doi.org/10.1016/s0020-7322(97)00017-2

    Keil, T.A. (1999) Morphology and development of the peripheral olfactory organs. In: Hansson, B.S. (Ed.), Insect olfaction. Springer, Berlin Heidelberg, pp. 5–47.
    https://doi.org/10.1007/978-3-662-07911-9_2

    Klots, A.B. (1931–1933) A generic revision of the Pieridae (Lepidoptera), together with a study of the male genitalia. Entomologica Americana, 12, 139–242.

    Koiwaya, S. (1996) Ten new species and twenty-four new subspecies of butterflies from China, with notes on the systematic positions of five taxa. In: Koiwaya, S. (Ed.), Studies of Chinese butterflies, 3, pp. 237–280.

    Lee, C.L. (1955) Some new species of Rhopalocera from China I. Description of a new pierid Leptidea serrata n. sp. Acta Entomologica Sinica, 5, 237–241. [in Chinese]

    Lelo, S. (2002) Variation in exogenous and endogenous (genitalia) characteristics of butterflies of the species Leptidea sinapis Linnaeus, 1758 (Pieridae, Dismorphiinae) within populations from the area around Sarajevo. Natura Croatica, 11 (3), 293–319.

    Linnaeus, C.V. (1758) Systema Naturae. Vol. 1. Edition X. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Holmiae Salvii, 824 pp.

    Llorente-Bousquets, J. (1984) Sinopsis Sistemática y Biogeográfica de los Dismorphiinae de México con especial referencia al género Enantia Huebner (Lepidoptera: Pieridae). Folia Entomologica Mexicana, 58, 1–207.

    Llorente-Bousquets, J. & Castro-Gerardino J. (2004) Subfamilia Dismorphiinae. In: LeCrom, J.F., Constantino, L.M., Llorente-Bousquets, J. & Salazar, J.A. (Eds.), Mariposas de Colombia Tomo 2: Pieridae. Ed. Carlec, Bogotá, pp. 14–16.

    Lorkovič, Z. (1993) Leptidea reali Reissinger, 1989 (= lorkovicii Réal 1988), a new European species (Lepid., Pieridae). Natura Croatica, 2 (1), 1–26.

    Lukhtanov, V.A., Dincă, V., Talavera, G. & Vila, R. (2011) Unprecedented within-species chromosome number cline in the Wood White butterfly Leptidea sinapis and its significance for karyotype evolution and speciation. BMC Evolutionary Biology, 11 (1), 109.
    https://doi.org/10.1186/1471-2148-11-109

    Maeki, J. (1958) On the cytotaxonomical relationships in Leptidea (Lepidoptera, Pieridae). Japan Journal of Genetics, 33 (9), 283–285.
    https://doi.org/10.1266/jjg.33.283

    Magallón, S., Gómez-Acevedo, S., Sánchez‐Reyes, L.L. & Hernández-Hernández, T. (2015) A metacalibrated time-tree documents the early rise of flowering plant phylogenetic diversity. New Phytologist, 207 (2), 437–453.
    https://doi.org/10.1111/nph.13264

    Martin, J.F., Gilles, A. & Descimon, H. (2003) Species concepts and sibling species: the case of Leptidea sinapis and Leptidea reali. In: Boggs C.L., Watt, W.B. & Ehrlich, P. (Eds.), Butterflies ecology and evolution: Taking flight. The University of Chicago Press, Chicago, pp. 459–476.

    Mazel, R. (2001a) Une sous-espèce asiatique de Leptidea reali Reissinger: L. reali yakovlevi ssp. nova (Lepidoptera, Pieridae, Dismorphiinae). Linneana Belgica, 18 (2), 103–104.

    Mazel, R. (2001b) Leptidea sinapis L., 1758—L. reali Reissinger, 1989, le point de la situation (Lepidoptea: Pieridae, Dismorphiinae). Linneana Belgica, 18, 199–202.

    Mazel, R. (2004) Leptidea descimoni species nova découverte au Kirghizistan (Lepidoptera, Pieridae). Linneana Belgica, 19 (5), 225–227.

    Mazel, R. (2005) Éléments de phylogénie dans le genre Leptidea Bilberg 1820 (Lepidoptera, Pieridae, Dismorphiinae). Revue l’Assoc roussillonaise d’Entomol, 14, 98–111.

    Mazel, R. & Eitschberger, U. (2002) Répartition géographique de Leptidea sinapis (L., 1758) et L. reali Reissinger, 1989 au nord de l’Europe, en Russie et dans quelqes pays d’Asie (Lepidoptera: Pieridae, Dismorphiinae). Linneana Belgica, 18 (8), 373–376.

    Mazel, R. & Leestmans, R. (1996) Relations biogéographiques, écologiques et taxonomiques entre Leptidea sinapis Linné et L. reali Reissinger en France, Belgique et régions limitrophes (Lepidoptera: Pieridae). Linneana Belgica, 15 (8), 317–331.

    Mazel, R. & Leestmans, R. (1999a) Séparation biométrique des Leptidea sinapis, L. morsei Fenton et L. reali Reissinger. Linneana Belgica, 17, 46–52.

    Mazel, R. & Leestmans, R. (1999b) Seconde contribution à l’étude des relations entre Leptidea sinapis Linné et L. reali Reissinger en France, Belgique et quelques autres contrées européennes (Lepidoptera: Pieridae). Linneana Belgica, 17, 155–168.

    Mazel, R. & Leestmans, R. (2003) Analyse et discussion de quelques types infraspécifiques dans le genre Leptidea Billberg, 1820 (Lepidoptera: Pieridae). I: types asiatiques. Linneana Belgica, 19 (4), 179–192.

    Mitter, K.T., Larsen, T.B., De Prins, W., De Prins, J., Collins, S., Vande Weghe, G., Sáfián, Sz., Zakharov, E.V., Hawthorne, D.J., Kawahara, A.Y. & Regier J.C. (2011) The butterfly subfamily Pseudopontiinae is not monobasic: marked genetic diversity and morphology reveal three new species of Pseudopontia (Lepidoptera: Pieridae). Systematic Entomology, 36 (1), 139–163.
    https://doi.org/10.1111/j.1365-3113.2010.00549.x

    Nelson, B., Hughes, M. & Bond, K. (2011) The distribution of Leptidea sinapis (Linnaeus, 1758) and L. reali Reissinger, 1989 (Lepidoptera: Pieridae) in Ireland. Entomologists Gazette, 62 (4), 213–233.

    Nelson, B., Hughes, M., Nash, R. & Warren, M. (2001) Leptidea reali Reissinger 1989 (Lep.: Pieridae): a butterfly new to Britain and Ireland. Entomologists Record and Journal of Variation, 113 (3), 97–102.

    Nieves-Uribe, S., Castro-Gerardino, J., Flores-Gallardo, A. & Llorente-Bousquets, J. (2016a) Corion en los géneros Anteos y Rhabdodryas (Pieridae: Coliadinae): su significado e implicaciones. Southwestern Entomologist, 41 (2), 485–504.
    https://doi.org/10.3958/059.041.0218

    Nieves-Uribe, S., Castro-Gerardino, J., Flores-Gallardo, A. & Llorente-Bousquets, J. (2016b) Microrretícula coriónica en los géneros Nathalis Boisduval y Krikogonia Reakirt (Lepidoptera: Pieridae): implicaciones taxonómicas y tendencias evolutivas. Southwestern Entomologist, 41 (3), 715–733.
    https://doi.org/10.3958/059.041.0314

    Nieves-Uribe, S., Castro-Gerardino, J., Flores-Gallardo, A. & Llorente-Bousquets, J. (2016c) Estudio del corion de tres especies del género Colias Fabricius, 1807 y Zerene cesonia cesonia (Stoll, 1790). Southwestern Entomologist, 41 (4), 1121–1141.
    https://doi.org/10.3958/059.041.0411

    Réal, P. (1988) Lépidoptères nouveaux, principalement jurassiens. Mémoires de Comité de Liaison pour les Recherches Écofaunistiques dans le Jura, 4, 1–28.

    Reissinger, E. (1989) Checkliste Pieridae Duponchel, 1835 der Westpalaearctis (Europa, Nordwestafrika, Kaukasus, Kleinasien). Atalanta, 20, 149–185.

    Reuter, E. (1896) Ueber die Palpen der Rhopaloceren. Ein Beitrag zur Erkenntniss der verwandtschaftlichen Beziehungen unter den Tagfaltern. Acta Societatia Scientiarum Fennicae, 22, 1–577.

    Sachanowicz, K. (2013) Separation possibilities and genital measurement variations in two cryptic species of European pierid butterflies, Leptidea juvernica Williams, 1946 and L. sinapis (Linnaeus, 1758). Zoology, 116 (4), 215–223.
    https://doi.org/10.1016/j.zool.2012.12.002

    Sachanowicz, K., Wower, A. & Buszko, J. (2011) Past and present distribution of the cryptic species Leptidea sinapis and L. reali (Lepidoptera: Pieridae) in Poland and its implications for the conservation of these butterflies. European Journal of Entomology, 108 (2), 235–242.
    https://doi.org/10.14411/eje.2011.032

    Sáfián, S. (2015) Behaviour and development of Pseudopontia gola Sáfián & Mitter, 2011 (Lepidoptera: Pieridae). SHILAP, Revista de Lepidopterología, 43 (169), 85–89.

    Schenk, O. (1903) Die antennalen Hautsinnesorgane einiger Lepidopteren und Hymenopteren. Zoologische Jahrbücher. Abteilung für Anatomie und Ontogenie der Tiere, 17, 573–618.
    https://doi.org/10.5962/bhl.part.4636

    Schmitz, O. (2007) Recent findings about the historic and current distribution of Leptidea sinapis (Linnaeus, 1758) and Leptidea reali Reissinger, 1989 (Lepidoptera, Pieridae) in the working area of the Rhenish-Westphalian Lepidopterologists. Entomologie Heute, 19, 181–195.

    Šíchová, J., Voleniková, A., Dincă, V., Nguyen, P., Vila, R., Sahara, K. & Marec, F. (2015) Dynamic karyotype evolution and unique sex determination systems in Leptidea wood white butterflies. BMC Evolutionary Biology, 15 (1), 1–16.
    https://doi.org/10.1186/s12862-015-0375-4

    Scott, J.A. (1984) The phylogeny of butterflies (Papilionoidea and Hesperioidea). Journal of Research on the Lepidoptera, 23 (4), 241–281.

    Sellier, R. (1974) Donnees documentaires sur l'ultrastructure des recepteurs sensoriels antennaires chez les Lepidopteres rhopaloceres; etude en microscopie electronique par balayage. Annales de la Societe Entomologique de France, 10, 917–937.

    Shields, V.D.C. (2008). Ultrastructure of insect sensilla. In: Capinera, J.L. (Ed.), Encyclopedia of Entomology. Springer, Dordrecht, pp. 4009–4023.
    https://doi.org/10.1007/978-1-4020-6359-6_2295

    Shields, V.D.C. & Hildebrand, J.G. (1999a) Fine structure of antennal sensilla of the female sphinx moth, Manduca sexta (Lepidoptera: Sphingidae). I. Trichoid and basiconic sensilla. Canadian Journal of Zoology, 77 (2), 290–301.
    https://doi.org/10.1139/z98-204

    Shields, V.D.C. & Hildebrand, J.G. (1999b) Fine structure of antennal sensilla of the female sphinx moth, Manduca sexta (Lepidoptera: Sphingidae). II. Auriculate, coeloconic, and styliform complex sensilla. Canadian Journal of Zoology, 77 (2), 302–313.
    https://doi.org/10.1139/z99-003

    Šíchová, J., Voleniková, A., Dincă, V., Nguyen, P., Vila, R., Sahara, K. & Marec, F. (2015) Dynamic karyotype evolution and unique sex determination systems in Leptidea wood white butterflies. BMC Evolutionary Biology, 15 (1), 1–16.
    https://doi.org/10.1186/s12862-015-0375-4

    Solovyev, V.I., Ilinsky, Y. & Kosterin, O.E. (2015) Genetic integrity of four species of Leptidea (Pieridae, Lepidoptera) as sampled in sympatry in West Siberia. Comparative cytogenetics, 9 (3), 299–324.
    https://doi.org/10.3897/compcytogen.v9i3.4636

    Steele, K.P. & Wojciechowski, M.F. (2003) Phylogenetic analyses of tribes Trifolieae and Vicieae, based on sequences of the plastid gene matK (Papilionoideae: Leguminosae). In: Klitgaard, B.B. & Bruneau, A. (Eds.), Advances in Legume Systematics. Part 10. Higher level systematics. Royal Botanic Gardens, Kew, pp. 355–370.

    Steinbrecht, R.A. (1999) Olfactory receptors. In: Eguchi, E. & Tominaga, Y. (Eds.), Atlas of Arthropod Sensory Receptors, Dynamic Morphology in Relation to Function. Springer-Verlag, Tokyo, pp. 155–76.

    Tennent, W.J. (1993) A critical note on the occurrence of Leptidea Billberg, 1820 species in North-West Africa (Lepidoptera: Pieridae). Phegea, 21 (4), 109–112.

    Tennent, J. (1996) The Butterflies of Morocco, Algeria, and Tunisia. Gem Publishing, Wallingford, 256 pp.

    Uchiyama, H., Awata, H., Kinoshita, M. & Arikawa, K. (2013) Rough eyes of the northeast-Asian wood white, Leptidea amurensis. Journal of Experimental Biology, 216 (18), 3414–3421.
    https://doi.org/10.1242/jeb.089169

    Verovnik, R. & Glogovčan, P. (2007) Morphological and molecular evidence of a possible hybrid zone of Leptidea sinapis and L. reali (Lepidoptera: Pieridae). European Journal of Entomology, 104 (4), 667–674.
    https://doi.org/10.14411/eje.2007.084

    Vila, R., Viader, S. & Jubany, J. (2003) Leptidea sinapis (Linnaeus, 1758) i L. reali (Reissinger 1988): dues ecpécies ‘‘bessones’’a Catalunya i Andorra (Lepidoptera: Pieridae). Bulleti de la Societat Catalana de Lepidopterologia, 90, 25–47.

    Wahlberg, N., Rota, J., Braby, M.F., Pierce, N.E. & Wheat, C.W. (2014) Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data. Zoologica Scripta, 43 (6), 641–650.
    https://doi.org/10.1111/zsc.12075

    Warren, M.S. (1985) The influence of shade on butterfly numbers in woodland rides, with special reference to the wood white Leptidea sinapis. Biological Conservation, 33 (2), 147–164.
    https://doi.org/10.1016/0006-3207(85)90101-6

    Warren, M.S., Pollard, E. & Bibby, T.J. (1986) Annual and long-term changes in a population of the wood white butterfly Leptidea sinapis. The Journal of Animal Ecology, 55, 707–719.
    https://doi.org/10.2307/4749

    Wiklund, C. (1977a) Oviposition, feeding and spatial separation of breeding and foraging habitats in a population of Leptidea sinapis (Lepidoptera). Oikos, 28, 56–68.
    https://doi.org/10.2307/3543323

    Wiklund, C. (1977b) Courtship behaviour in relation to female monogamy in Leptidea sinapis (Lepidoptera). Oikos, 29, 275–283.
    https://doi.org/10.2307/3543614

    Wiklund, C. (1984) Egg-laying patterns in butterflies in relation to their phenology and the visual apparency and abundance of their host plants. Oecologia, 63 (1), 23–29.
    https://doi.org/10.1007/bf00379780

    Wojciechowski, M.F., Sanderson, M.J., Steele, K.P. & Liston, A. (2000) Molecular phylogeny of the ‘Temperate’ herbaceous tribes of Papilionid legumes: a super tree approach. In: Herendeen, P.S. & Bruneau, A. (Eds.), Advances in Legume Systematics. Vol. 9. Royal Botanic Gardens, Kew, pp. 277–298.