Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2018-06-06
Page range: 201–246
Abstract views: 217
PDF downloaded: 3

Chorionic sculpture of eggs in the subfamily Dismorphiinae (Lepidoptera: Papilionoidea: Pieridae)

Museo de Zoología (Entomología), Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, C.P. 04510, CDMX, México. Smithsonian Institution, PO Box 37012, Washington, DC. 20013-7012 USA.
Museo de Zoología (Entomología), Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, C.P. 04510, CDMX, México.
Museo de Zoología (Entomología), Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, C.P. 04510, CDMX, México.
Museo de Zoología (Entomología), Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, C.P. 04510, CDMX, México.
Smithsonian Institution, PO Box 37012, Washington, DC. 20013-7012 USA.
Lepidoptera Papilionoidea Pieridae

Abstract

This is the second exploration, comparison, and analysis of the chorion of species (45 sspp.) of the subfamily Dismorphiinae (Pieridae). This study includes nearly 50% of the species of the subfamily, including six of the seven genera in its two subtribes: Leptidea (Leptideini), Enantia, Pseudopieris, Lieinix, Moschoneura, and Dismorphia (Dismorphiini). The material studied originates from more than three dozen localities in six different countries on three continents (America, Asia, and Europe) and two biogeographical regions, the Palearctic and Neotropical, over the last 20 years. We have corrected and added information regarding several morphological aspects of the chorion. The precision of the citriform configuration and the elimination of the meloniform shape in the egg of Dismorphiini were determined with detailed observations on the maturation of the chorion in the ovarioles where each stage appears in a linear sequence. We discerned that the meloniform states correspond to incompletely differentiated or immature eggs. This was confirmed by the study of new samples of Dismorphia amphione, D. eunoe, and D. lewyi. The chorion of Dismorphiinae is basically plesiomorphic with respect to those of Coliadinae and Pierinae because it lacks several typical synapomorphies of these subfamilies, such as the presence of micro-grid and/or perimicropylar and apex differentiation, respectively. The eggs of each Dismorphiinae genus can be diagnosed by a combination of chorionic features, although sometimes by one or more plesiomorphies or apomorphies in each genus, with respect to the form or character states in axes, ribs, and poles in the grid of the three regions of the egg - two polar regions and one equatorial (basal, medial, and apical). Leptidea and Enantia show the most generalized grid pattern; however, two genera retain several plesiomorphies with respect to the undifferentiated axes or a small number of short axes (Pseudopieris), as well as many equidistant ribs (Lieinix). The chorionic grid of Moschoneura, although practically lacking short axes, shows the fewest number of axes in the entire subfamily (eight aligns it with Pseudopieris). The chorionic grid in Dismorphia is highly diverse, as it shows the most derived states; however, it comprises symplesiomorphies or atavisms in two groups of species, which aligns them closer to Lieinix or Pseudopieris, but we do not take them into account in some cases where they are convergences or structural parallelisms. It seems that the combination of the shape and its length:width ratio is correlated with the alar configuration (design, sexual dimorphism, and coloring patterns) and separates three groups of species in Dismorphia, and often correlates with the number of ribs. This also coincides with the Batesian participation in the number of mimetic complexes in which a subgroup of species and their stenoecy are integrated within the primary forests. Finally, two schemes are presented that synthesize and illustrate the changes or progression of the form and chorionic grid in the genera of the subfamily.

 

References

  1. Ackery, P.R., de Jong, R. & Vane-Wright, R.I. (1999) The butterflies: Hedyloidea, Hesperioidea and Papilionoidea. In: Kristensen, N.P. (Ed.), Handbook of Zoology. Vol. IV. Arthropoda: Insecta. Part 35. Lepidoptera, Moths and Butterflies. Vol. 1. Evolution, Systematics and Biogeography. Walter de Gruyter, NY, 263 pp.

    Aiello, A. (1980) Life history of Dismorphia amphione beroe (Lepidoptera: Pieridae: Dismorphiinae) in Panama. Psyche, 87 (3–4), 171–176.
    https://doi.org/10.1155/1980/38348

    Braby, M.F. (2005) Provisional checklist of genera of the Pieridae (Lepidoptera: Papilionidae). Zootaxa, 832 (1), 1–16.
    https://doi.org/10.11646/zootaxa.832.1.1

    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

    Chapman, R.F. (1998) The Insects, Structure and Function. 4th Edition. Cambridge University Press, Cambridge, 771 pp.
    https://doi.org/10.1017/CBO9780511818202

    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

    Dolinskaya, I.V. (2010) The chorionic sculpture in eggs of some Hadeninae (Lepidoptera, Noctuidae) from Ukraine. Ukrainska Entomofaunistyka, 1 (3), 3–15.

    Dolinskaya, I.V. (2011) The chorionic sculpture of the eggs of some Xyleninae (Lepidoptera, Noctuidae). Vestnik Zoologii, 45 (1), 7–22.
    https://doi.org/10.2478/v10058–011–0002–9

    Dolinskaya, I.V. & Geryak, Y.N. (2010) The chorionic sculpture of the eggs of some Noctuinae (Lepidoptera, Noctuidae) from Ukraine. Vestnik Zoologii, 44 (5), 421–432.
    https://doi.org/10.2478/v10058–010–0028–4

    Dolinskaya, I.V. & Ponomarenko, M.G. (2013) The chorionic sculpture in eggs of some Noctuidae (Lepidoptera). Vestnik Zoologii, 47 (5), 33–41.
    https://doi.org/10.2478/vzoo–2013–0046

    Döring, E.K.H. (1955) Zur Morphologie der Schmetterlingseier. Akademie-Verlag, Berlín, 154 pp., 61 pls.

    Downey, J.C. & Allyn, A.C. (1980) Eggs of Riodinidae. Journal of the Lepidopterists' Society, 34 (2), 133–145.

    Downey, J.C. & Allyn, A.C. (1981) Chorionic sculpturing in eggs of Lycaenidae. Part I. Bulletin of the Allyn Museum, 61, 1–29.

    Eitschberger, U. & Strohle, M. (1990) Zehnte Ergänzung zu “Systematische Unterschungen am Pieris-napi-bryoniae Komplex (s.l.)”. Aufzucht und Beschreibung der Praimaginalstadien von Pieris bryoniae adalwinda Fruhstorfer, 1909 und der Phaenotyp der Imagines (Lepidoptera, Pieridae). Atalanta, 21, 239–252.

    Ehrlich, P.R. (1958) The comparative morphology, phylogeny and higher classification of the butterflies (Lepidoptera: Papilionoidea). The University of Kansas Science Bulletin, 39 (8), 305–370.

    Fehrenbach, H. (2003) Eggs. In: Kristensen, N.P. (Ed.), Handbook of Zoology: Vol. IV. Arthropoda: Insecta, Part 36: Lepidoptera, Moths and Butterflies. Vol. 2. Morphology, Physiology and Development. Walter de Gruyter, New York, pp. 469–493.

    Ford, E.B. (1945) Butterflies. Collins, London, 368 pp.

    García-Barros, E. (1992) Evidence for geographic variation of egg size and fecundity in a Satyrine butterfly, Hypparchia senile (L.) (Lepidoptera, Nymphalidae-Satyrinae). Graellsia, 48, 45–52.

    García-Barros, E. (2000a) Body size, egg size, and their interspecific relationships with the ecological and life history traits in butterflies (Lepidoptera: Papilionoidea, Hesperioidea). Biological Journal of the Linnean Society, 70 (2), 251–284.
    https://doi.org/10.1111/j.1095–8312.2000.tb00210.x

    García-Barros, E. (2000b) Egg size in butterflies (Lepidoptera: Papilionoidea and Hesperiidae): a summary of data. The Journal of Research on the Lepidoptera, 35, 90–136.

    García-Barros, E. & Martín, J. (1995) The eggs of the European satyrine butterflies (Nymphalidae): external morphology and its use in systematics. Zoological Journal of the Linnean Society, 115 (1), 73–115.
    https://doi.org/10.1111/j.1096–3642.1995.tb02324.x

    García-Barros, E. & Munguira, M.L. (1997) Uncertain branch lengths, taxonomic sampling error, and the egg to body size allometry in temperate butterflies (Lepidoptera). Biological Journal of the Linnean Society, 61 (2), 201–221.
    https://doi.org/10.1111/j.1095–8312.1997.tb01787.x

    Häuser, C.L. (1993) Die inheren weiblichen Genitalorgane der Tagfalter (Rhopalocera): Vergleichende Morphologie und philogenetische Interpretarion (Insecta: Lepidoptera). Zoologische Jahrbücher. Abbteilung für Systematik, Okölogie und Geographie der Tiere, 120 (4), 389–439.

    Hernández, J.M. (1991) Estudio de los caracteres del huevo en diversos Cerambycidae Ibéricos y su interés taxonómico (Coleoptera). Graellsia, 47, 49–59.

    Hernández-Mejía, B.C., Flores-Gallardo, A. & Llorente-Bousquets, J. (2013) Comparación morfológica del corion de especies de los géneros Pieriballia, Itaballia y Perrhybris (Lepidoptera: Pieridae: Pierinae), y sus implicaciones filogenéticas. Southwestern Entomologist, 38 (2), 275–292.
    https://doi.org/10.3958/059.038.0211

    Hernández-Mejía, B.C., Flores-Gallardo, A. & Llorente-Bousquets, J. (2014a) Morfología del corion en especies de los géneros Ascia y Ganyra y su comparación con otros géneros próximos de Pierinae (Lepidoptera: Pieridae). Southwestern Entomologist, 39 (1), 119–134.
    https://doi.org/10.3958/059.039.0112

    Hernández-Mejía, B.C., Flores-Gallardo, A. & Llorente-Bousquets, J. (2014b) Morfología del Corion en la Subfamilia Coliadinae (Lepidoptera: Pieridae). Southwestern Entomologist, 39 (4), 853–886.
    https://doi.org/10.3958/059.039.0416

    Hernández-Mejía, C., Flores-Gallardo, A. & Llorente-Bousquets J. (2015) Morfología del Corion en Leptophobia (Lepidoptera: Pieridae) e Importancia Taxonómica. Southwestern Entomologist, 40 (2), 351–368.
    https://doi.org/10.3958/059.040.0210

    Hernández-Roldán, J.L., Munguira, M.L., Wagner, W. & Vila, R. (2012) Comparative analysis and taxonomic use of the morphology of immature stages and natural history traits in European species of Pyrgus Hübner (Lepidoptera: Hesperiidae, Pyrginae). Zootaxa, 3470, 1–71.

    Hinton, H.E. (1981) Biology of Insect Eggs. Vol. I–III. Pergamon, Oxford, 1125 pp.

    Howe, W.H. (1975) The Butterflies of North America. Doubleday & Company, Garden City, New York, 633 pp., 97 pls.

    Jasso-Martínez, J.M., Castañeda-Sortibrán, A.N., Pozo, C., García–Sandoval, R., Prado, B.R., Luis-Martínez, A., Llorente-Bousquets, J. & Rodríguez-Arnaiz, R. (2016) The Enantia Jethys Complex 1: Insights from COI confirm the species complex and reveal a new potential cryptic species. Southwestern Entomologist, 41 (4), 1005–1020.
    https://doi.org/10.3958/059.041.0401

    Kambysellis, M.P., Margaritis, L. & Craddock, E.M. (1999) Eggs coverings, insects. In: Knobil, E. & Neil, J. (Eds.), Encyclopedia of Reproduction. Vol. I. Academic Press, New York, pp. 971–990.

    Kristensen, N.P. (1975) Remarks on the family–level phylogeny of butterflies (Insecta, Lepidoptera, Rhopalocera). Journal of Zoological Systematics and Evolutionary Research, 14 (1), 25–33.
    https://doi.org/10.1111/j.1439–0469.1976.tb00515.x

    Kristensen, N.P. (2003) Reproductive organs. In: Kristensen, N.P. (Ed.), Handbook of Zoology. Vol. IV Arthropoda: Insecta. Part 36. Lepidoptera, Moths and Butterflies. Vol. 2. Morphology, physiology and development. Walter de Gruyter, New York, pp. 427–447.

    Küppers, H. (1996) Atlas de los colores, más de 5500 matices con su caracterización y las instrucciones para su mezcla. Blume, Barcelona, 161 pp.

    Kumar, V., Kariappa, B.K., Babu, A.M. & Dandin, S.B. (2007) Surface ultrastructure of the egg chorion of Eri silkworm Samia ricini (Donovan) (Lepidoptera: Saturniidae). Journal of Entomology, 4 (2), 68–81.
    https://doi.org/10.3923/je.2007.68.81

    Lamas, G. (1979) Los Dismorphiinae (Pieridae) de México, América Central, y las Antillas. Revista de la Sociedad Mexicana de Lepidopterología, 5 (1), 3–37.

    Llorente-Bousquets, J. (1984) Sinopsis sistemática y biogeográfica de los Dismorphiinae de México con especial referencia al género Enantia Hübner (Lepidoptera: Pieridae). Folia Entomológica Mexicana, 58, 3–206.

    Llorente-Bousquets, J. & Castro-Gerardino, J. (2004) Subfamilia Dismorphiinae. In: LeCrom, J.F., Llorente-Bousquets, J., Constantino, L.M. & Salazar, J.A. (Eds.), Mariposas de Colombia. Tomo 2. Pieridae. Carlec, Bogotá, pp. 14–15.

    Llorente-Bousquets, J. & Castro-Gerardino, J. (2007) Estudios en sistemática de Dismorphiini (Lepidoptera: Pieridae) I. Morfología de huevos y su importancia taxonómica. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 31 (118), 145–164.

    Llorente-Bousquets, J. & Castro-Gerardino, J. (2008) Morfología del corion en Dismorphiini (Lepidoptera: Pieridae). In: Llorente-Bousquets, J. & Lanteri, A. (Eds.), Contribuciones Taxonómicas en Órdenes de Insectos Hiperdiversos. Las Prensas de Ciencias, UNAM, México, D.F., pp. 72–90.

    Llorente-Bousquets, J. & Castro-Gerardino, D.J. (2018) Antennal ultrastructure of Leptidea (Pieridae, Dismorphiinae, Leptideini) and its taxonomic implications. Zootaxa, 4402 (3), 401–442.
    https://doi.org/10.11646/zootaxa.4402.3.1

    Llorente-Bousquets, J. & Garcés, A.R. (1983) Notas sobre Dismorphia amphiona lupita Lamas (Lepidoptera: Pieridae) y observaciones sobre algunos complejos miméticos en México. Revista de la Sociedad Mexicana de Lepidopterología, 8, 27–39.

    Llorente-Bousquets, J. & LeCrom, J.F. (2013) Descripción de dos nuevos Dismorphiini (Lepidoptera: Pieridae) con reflexiones del endemismo en la Sierra Nevada de Santa Marta, Colombia. Revista Colombiana de Entomología, 39 (2), 276–280.

    Llorente-Bousquets, J., Castro-Gerardino, D.J. & LeCrom, J.F. (2016) Estudio del género Moschoneura Butler, 1870 y una nueva subespecie de Moschoneura ela (Lepidoptera: Pieridae, Dismorphiinae). Southwestern Entomologist, 41 (4), 1143–1160.
    https://doi.org/10.3958/059.041.0412

    Margaritis, L.H. (1985) Structure and physiology of the eggshell. In: Kerkut, G.A. & Gilbert, L.I. (Eds.), Comprehensive Insect Phisiology Biochemistry and Pharmacology. Vol. I. Pergamon press, New York, pp. 1583–230.

    Margaritis, L.H. & Mazzini, M. (1998) Structure of the egg. In: Harrison, F.W. & Locke, M. (Eds.), Microscopic Anatomy of Invertebrates. Vol. 11b. Insecta. Wiley-Liss, New York, pp. 995–1037.

    Mazur, G.D., Regier, J.C. & Kafatos, F.C. (1982) Order and defects in the silkmoth chorion, a biological analogue of a cholesteric liquid crystal. In: Akai, H. & King, R.C. (Eds.), Insect ultrastructure. Vol. I. Plenum Press, New York, pp. 150–185.
    https://doi.org/10.1007/978-1-4615-7266-4_6

    Munguira, M.L., Martín, J., García-Barros, E., Shahbazian, G. & Cancela, J.P. (2015) Morphology and morphometry of Lycaenid eggs (Lepidoptera: Lycaenidae). Zootaxa, 3937 (2), 201–247.
    https://doi.org/10.11646/zootaxa.3937.2.1

    Nakasuji, F. (1987) Egg size of skippers (Lepidoptera: Hesperiidae) in relation to their host specificity and to leaf toughness of host plants. Ecological Research, 2 (2), 175–183.
    https://doi.org/10.1007/BF02346925

    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 Kricogonia Reakirt (Lepidoptera: Pieridae): implicaciones taxonómicas y tendencias evolutivas. Southwestern Entomologist, 41 (3), 715–734.
    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

    Nieves-Uribe, S., Flores-Gallardo, A., Hernández-Mejía, B.C. & Llorente-Bousquets, J. (2015) Exploración morfológica del corion en Biblidinae (Lepidoptera: Nymphalidae): aspectos filogenéticos y clasificatorios. Southwestern Entomologist, 40 (3), 589–648.
    https://doi.org/10.3958/059.040.0318

    Nieves-Uribe, S., Flores-Gallardo, A. & Llorente-Bousquets, J. (2016d) Morfología coriónica de once especies de Biblidinae de México y Colombia: un examen de predicciones. Southwestern Entomologist, 41 (2), 505–532.
    https://doi.org/10.3958/059.041.0219

    Nieves-Uribe, S., Flores-Gallardo, A. & Llorente-Bousquets, J. (2017a) Corion de Ectima thecla thecla (Fabricius, 1796) y su Comparación con el Género Hamadryas. Southwestern Entomologist, 42 (3), 851–864.
    https://doi.org/10.3958/059.042.0324

    Nieves-Uribe, S., Flores-Gallardo, A. & Llorente-Bousquets, J. (2017b) Corion de Orophila cardases campaspe (Hewitson, 1869) y caracterización coriónica de la subtribu Callicorina. Southwestern Entomologist, 42 (3), 865–878.
    https://doi.org/10.3958/059.042.0325

    Papageorgis, C. (1975) Mimicry in Neotropical butterflies. American Scientist, 63 (5), 522–532.

    Peterson, A. (1948) Larvae of insects. An introduction to Neartic species. Part I. Lepidoptera and plant infesting Hymenoptera. 3th Edition. Published by the author, Ohio State University, Columbus, OH, 315 pp.

    Peterson, A. (1960) Photographing eggs of insects. The Florida Entomologist, 43 (1), 1–7.
    https://doi.org/10.2307/3492514

    Peterson, A. (1961) Some types of eggs deposited by moths, Heterocera– Lepidoptera. The Florida Entomologist, 44 (3), 107–114.
    https://doi.org/10.2307/3492966

    Peterson, A. (1962) Some eggs of moths among the Geometridae—Lepidoptera. The Florida Entomologist, 45 (3), 109–119.
    https://doi.org/10.2307/3492561

    Peterson, A. (1963a) Some eggs of moths among the Amatidae, Arctiidae, and Notodontidae—Lepidoptera. The Florida Entomologist, 46 (2), 169–182.
    https://doi.org/10.2307/3493627

    Peterson, A. (1963b) Egg types among moths of the Pyralidae and Phycitidae— Lepidoptera. The Florida Entomologist, 46 (Supplement), 1–14.
    https://doi.org/10.2307/3493354

    Peterson, A. (1964) Egg types among moths of the Noctuidae (Lepidoptera). The Florida Entomologist, 47 (2), 71–91.
    https://doi.org/10.2307/3493280

    Peterson, A. (1965a) Some eggs of moths among the Olethreutidae and Tortricidae (Lepidoptera). The Florida Entomologist, 48 (1), 1–8.
    https://doi.org/10.2307/3493515

    Peterson, A. (1965b) Some eggs of moths among the Sphingidae, Saturniidae, and Citheroniidae (Lepidoptera). The Florida Entomologist, 48 (4), 213–219.
    https://doi.org/10.2307/3493773

    Peterson, A. (1966) Some eggs of moths among the Liparidae, Lasiocampidae, and Lacosomidae (Lepidoptera). The Florida Entomologist, 49 (1), 35–42.
    https://doi.org/10.2307/3493314

    Peterson, A. (1967a) Some eggs of moths from several families of Microlepidoptera. The Florida Entomologist, 50 (2), 125–132.
    https://doi.org/10.2307/3493621

    Peterson, A. (1967b) Eggs of moths among the Ethmiidae, Acrolophidae and Hepialidae—Microlepidoptera. The Florida Entomologist, 50 (3), 181–183.
    https://doi.org/10.2307/3493299

    Peterson, A. (1968) Eggs of moths from additional species of Geometridae— Lepidoptera. The Florida Entomologist, 51 (2), 83–94.
    https://doi.org/10.2307/3493606

    Peterson, A. (1970) Eggs from miscellaneous species of Rhopalocera— Lepidoptera. The Florida Entomologist, 53 (2), 65–71.
    https://doi.org/10.2307/3493448

    Rezende, G.L., Vargas, H.C.M., Moussian, B. & Cohen, E. (2016) Composite eggshell matrices: Chorionic layers and sub-chorionic cuticular envelopes. In: Cohen, E., Moussian, B. (Eds.), Extracellular composite matrices in arthropods. Springer International Publishing, Cham, pp. 325–366.
    https://doi.org/10.1007/978-3-319-40740-1_9

    Richards, W. & Davies, R.G. (Eds). (1977) Imms' general textbook of entomology. Vol.I. Structure, Physiology and Development. Chapman and Hall, London, 418 pp.

    Sariot, M.G.M. (2016) Descripción del huevo y estados larvarios de Elphinstonia bazae (Fabiano, 1993) (Lepidoptera: Pieridae). Boletin de la Sociedad Entomológica Aragonesa, 59 (2), 207–211.

    Skudlik, J., Poprawa, I. & Rost, M.M. (2005) The egg capsule of Spodoptera exiqua Hübner, 1808 (Insecta, Lepidoptera, Noctuidae): morphology and ultrastructure. Zoologica Poloniae, 50 (1–4), 25–31.

    Sourakov, A. (2011) Report of partial bilateral gynandromorph of Dismorphia spio with notes on adult sexual dimorphism and illustrations of immature stages. Journal of the Lepidopterists’ Society, 65 (3), 196–199.
    https://doi.org/10.18473/lepi.v65i3.a10

    Thomson, G. (1992) Egg surface morphology of Manioline butterflies (Lepidoptera, Nymphalidae, Satyrinae). Atalanta, 23 (1/2), 195–214.

    Torres-Bauzá, J.A. (1991) Biología de Dismorphia spio (Godart) en Puerto Rico (Lepidoptera: Pieridae: Dismorphiinae). Caribbean Journal of Science, 27 (1–2), 35–45.

    Trougakos, I.P. & Margaritis, L.H. (2008). Novel morphological and physiological aspects of insect eggs. In: Hilker, M. & Meiners, T. (Eds.), Chemoecology of Insect Eggs and Egg Deposition. Blackwell Publishing Company, Berlin, pp. 3–36.

    Vane-Wright, R.I. (2003) Evidence and identity in butterfly systematics. In: Boggs, C.L., Watt, W.B. & Ehrlich, P.R. (Eds.), Butterflies: ecology and evolution taking flight. Chicago University Press, Chicago, IL, pp. 477–514.

    Walhberg, N., Rota, J., Bravy, 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

    Wheeler, D.E. (2009) Egg Coverings. In: Resh, V.H. & Cardé, R.T. (Eds.), Encyclopedia of Insects. Academic Press, Amsterdam, pp. 312–313.
    https://doi.org/10.1016/B978-0-12-374144-8.00093-X

    Winhard, W. (1996) Konvergente Farbmusterentwicklungen bei Tagfaltern. Spixiana, 21 (Supplement), 1–192.

    Winhard, W. (2016) Was macht Schmetterlinge ähnlich, wenn nicht verwandtschaft? Erläutert an Beispielen aus Südamerika, hauptsächtlich Kolumbien und Ecuador mit Neubeschreibungen einer Art und von 17 Unterarten (Lepidoptera, Pieridae, Nymphalidae). Atalanta, 47 (3/4), 290–319.

    Young, A.M. (1972) Notes on the life cycle and natural history of Dismorphia virgo (Lepidoptera: Pieridae: Dismorphinae) in Costa Rica. Psyche, 79 (3), 165–178.
    https://doi.org/10.1155/1972/82529