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Constantino, L. M. ., & Constantino, S. . (2020). Introgresión genética y especiación de Heliconius hecalesia (Hewitson [1854]) (Lepidoptera: Heliconiinae) en zonas de hibridación en el occidente de Colombia. Boletín Científico Centro De Museos Museo De Historia Natural, 24(2), 206–230. https://doi.org/10.17151/bccm.2020.24.2.14
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Luis Miguel Constantino
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Sandra Constantino
Resumen
aclarar el estatus taxonómico de Heliconius gynaesia y H. longarena dentro del grupo de “hecalesia” y conocer la delimitación genética y geográfica de las subespecies. Metodología: se realizaron observaciones directas en campo en las áreas de distribución de las diferentes subespecies de H. hecalesia en Colombia. Para comparar los diferentes morfotipos se realizaron cruces genéticos en condiciones de invernadero entre las poblaciones de H. hecalesia hecalesia del Magdalena medio, con las poblaciones de H. hecalesia longarena de la región del Chocó biogeográfico en el occidente de Colombia. Resultado: se presentan y discuten los cruces genéticos de los diferentes fenotipos obtenidos de un mismo cruce parental y retrocruce, del cual fue posible reconstruir las formas gynaesia y ernestus, que son consistentes con el patrón de coloración a los especímenes tipo. Todos los fenotipos estudiados están involucrados en un nuevo complejo mimético con Tithorea tarricina y Neruda godmani, el cual es descrito en este artículo. El aislamiento reproductivo de las poblaciones del suroccidente de Colombia que se manifiesta mediante esterilidad hibrida e inviabilidad en los cruces artificiales, proporciona una fuerte evidencia y soporte para el reconocimiento de H. longarena n. comb como una buena especie como parte de este estudio. Se coloca a H. hecalesia gynaesia en sinonimia n. syn con H. longarena por tratarse de un híbrido natural.
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Citas
Ackery, P.R. y Smiles, R. (1976). An illustrated list of the type-specimens of the Heliconiinae (Lepidoptera: Nymphalidae) in the British Museum (Natural History) Bulletin of the British Museum of Natural History, 32 (5), 173-214 + 39 pls.
Arias, C.F., Muñoz, A.G., Jiggins, C.D., Mavarez, J., Bermingham, E. y Linares, M. (2008) A bimodal hybrid zone provides evidence for incipient speciation in Heliconius butterflies. Molecular Ecology, 17, 4699-4712.
Audesirk, T. y Audesirk, G. (1996). El origen de las especies. Pp. 46-58. En: Audesirk, T. & Audesirk, G. (Ed). Evolución y ecología. Cuarta Edición. Prentice Hall Hispanoamericana S.A. México. 407 p.
Barton, N.H. y Hewitt, G.M. (1989). Adaptation, speciation and hybrid zones. Nature, 341, 497-503.
Brower, A.V. (1996). A new mimetic species of Heliconius (Lepidoptera: Nymphalidae), from southeastern Colombia revealed by cladistic analysis of mithochondrial DNA sequences. Zoological Journal of the Linnean Society, 116, 317-332.
Brower, A.V. (2011). Hybrid speciation in Heliconius butterflies? A review and critique of the evidence. Genetica, 139, 589-609.
Brown, K.S. y Benson, W. (1975). West Colombian biogeography. Notes on Heliconius hecalesia and H. sapho (Nymphalidae). Journal of the lepidopterists´ Society, 29 (4), 199-212.
Brown, K.S. (1975). Geographical patterns of evolution in Neotropical Lepidoptera. Systematics and derivation of known and new Heliconiini (Nymphalidae: Nymphalinae). Journal of Entomology
(B), 44 (3), 201-242.
Brown, K.S. (1981). The biology of Heliconius and related genera. Ann. Rev. Entomol., 26, 427-456.
Brown, K.S., Emmel, T.C., Eliazar, P.J. y Suomalainen, E. (1992). Evolutionary patterns in chromosome numbers in Neotropical Lepidoptera. I. Chromosomes of the Heliconiini. Hereditas, 117, 109–125.
Constantino, L.M. y Salazar, J. (1998). Natural hybridization of Heliconius cydno Doubleday from Western Colombia (Lepidoptera: Nymphalidae: Heliconiinae). Bol. Cient. Mus. Hist. Nat. U. Caldas, 2, 41-45.
Constantino, L.M., Gil, Z.N. y Corredor, G. (2005). Estudio del polimorfismo cromático en Laparus doris obscurus y determinación de la frecuencia fenotípica mediante cruzamientos intraespecíficos de tres poblaciones en el departamento del Valle y Caldas, Colombia (Nymphalidae: Heliconinae). Bol. Cient. Mus. Hist. Nat Universidad de Caldas,9, 222-237.
Constantino, L.M. y Salazar, J.A. (2010). A review of the Philaethria dido species complex (Lepidoptera. Nymphalidae: Heliconiinae) and description of three new sibling species from Colombia and Venezuela. Zootaxa, 2720, 1- 27.
Constantino, L.M. (2006). Redescubrimiento de Heliconius weymeri submarginatus, 98 años después de su aparente extinción y estudio de su complejo mimético con Heliconius hecubacrispus en el occidente de Colombia. Bol. Cient. Mus. Hist. Nat. Universidad de Caldas, 10, 138-197.
Constantino, L.M. (2016). Análisis morfológicos, moleculares y biogeográficos en la validación de nuevas especies y resolución de problemas taxonómicos en Lepidoptera. Memorias 43 Congreso Sociedad Colombiana de Entomología, Socolen. Manizales, 43, 209-228.
Dasmahapatra, K.K., Elias, M., Hill, R.I., Hoffman, J. y Mallet, J. (2010). Mitochondrial DNA barcoding detects some species that are real, and some that are not. Molecular Ecology Resources, 10, 264-273.
Dobzhansky, T. (1937). Genetics and the origin of species. Columbia University Press, New York.
Ebach, M.C. y Holdrege, C. (2005). DNA barcoding is no substitute for taxonomy. Nature, 434, 697-697.
Edelman, N.B., Frandsen, P.B., Miyagi, M., Clavijo, B., Davey, J., Dikow, R.B., García-Accinelli, G., Van Belleghem, S.M., Patterson, N., Neafsey, D. E., Challis, R., Kumar, S., Moreira, G.R.P., Salazar, C., Chouteau, M., Counterman, B.A., Papa, R., Blaxter, M., Reed, R.D., ... Mallet, J. (2019). Genomic architecture and introgression shape a butterfly radiation. Science, 366 (6465), 594-599.
Elias, M., Hill, R.I., Willmott, K.R., Dasmahapatra, K.K., Brower, A. V., Mallet, J. y Jiggins, C.D. (2007). Limited performance of DNA barcoding in a diverse community of tropical butterflies. Proceedings of the Royal Society of London B: Biological Sciences, 274 (1627), 2881-2889.
Gilbert, L.E. (2003). Adaptive novelty through introgression in Heliconius wing patterns: evidence for a shared genetic "toolbox" from synthetic hybrid zones and a theory of diversification. Ecology and Evolution Taking Flight: Butterflies as Model Systems. Edited by: Boggs C.L., Chicago: University of Chicago Press, 281-318.
Giraldo, C.E. (2015). La taxonomía integrativa en la resolución de problemas taxonómicos complejos en insectos: un caso de estudio en mariposas diurnas neotropicales. Libro de Memorias, 42 Congreso Sociedad Colombiana de Entomología, Medellín.,42, 142- 162.
Hendry, A.P. (2009). Speciation. Nature, 458, 162-164.
Jiggins, C.D., McMillan, W.O., Neukirchen, W. y Mallet, J. (1996). What can hybrid zones tell us about speciation? The case of Heliconius erato and H. himera (Lepidoptera: Nymphalidae). Biol J Linn Soc., 59, 221-242.
Jiggins, C.D. y McMillan, W.O. (1997). The genetics basis of an adaptive radiation: warning colour in two Heliconius species. Proc. R. Soc. Lond. B., 264, 1167-1175.
Jiggins, C.D. y Mallet, J. (2000). Bimodal hybrid zones and speciation. Trends Ecol Evol., , 15, 250-255.
Jiggins, C.D., Naisbit, R.E., Coe, R.L. y Mallet, J. (2001). Reproductive isolation caused by colour pattern mimicry. Nature, 411, 302-305.
Jiggins, C.D., Salazar, C., Linares, M. y Mavarez, J. (2008). Hybrid trait speciation and Heliconius butterflies. Speciation special issue of Phil. Trans. Royal. Soc., 363, 3047-3054.
Kozak, K.M., Wahlberg, N., Neild, A.F., Dasmahapatra, K.K., Mallet, J. y Jiggins, C.D. (2015). Multilocus species trees show the recent adaptive radiation of the mimetic Heliconius butterflies. Systematic Biology 64, 505-524.
Kronforst., M.R., Young, L.G., Blume, L. M. y Gilbert, L.E. (2006). Multilocus analysis of admixture and introgression among hybridizing Heliconius butterflies. Evolution, 60, 1254-1268.
Lamas, G. (2004). Heliconiinae pp. 261-274. In. Lamas, G. (Ed.), Atlas of Neotropical Lepidoptera, Check list: Part 4A, HesperoideaPapilionoidea. Association for Tropical Lepidoptera, Gainesville. 439 pp.
Linares, M. (1996). The genetics of the mimetic coloration in the butterfly Heliconius cydno weymeri. Heredity, 87, 142-149.
Mallet, J. y Barton, N. H. (1989). Strong natural selection in a warning colour hybrid zone. Evolution, 43, 421-431.
Mallet, J. (1993). Speciation, radiation, and color pattern evolution in Heliconius butterflies: evidence from hybrid zones. Hybrid Zones and the Evolutionary Process. Edited by: Harrison RG. New York: Oxford University Press, 226-260.
Mallet, J. (1995). A species definition for the Modern Synthesis. Tree, 10, 294-298.
Mallet, J. y Gilbert, L.E., 1995.- Why are there so many mimicry rings? Correlations between habitat, behaviour and mimicry in Heliconius butterflies. Biol J Linn Soc., 55, 159-180.
Mallet, J. (1999). Causes and consequences of a lack of coevolution in Müllerian mimicry. Evolutionary Ecology, 13, 777-806.
Mallet, J. (2007). Hybrid speciation. Reviews. Nature, 446, 279-283.
Mallet, J. (2013). Species, concept of. Encyclopedia of Biodiversity. Volume 6, 679-691.
Mallet, J. y Willmott, K. (2003). Taxonomy: renaissance or Tower of Babel? Trends Ecol. Evol., 18, 57-59.
Mallet, J., Beltran, M., Neukirchen, W. y Linares, M. (2007). Natural hybridization in Heliconiine butterflies: the species boundary as a continuum. BMC Evolutionary Biology. 7,1-16.
Martin, S.H., Dasmahapatra, K.K., Nadeau, N.J., Salazar, C., Walters, J.R., Simpson, F., Blaxter, M., Manica, A., Mallet, J. y Jiggins, C.D. (2013). Genome-wide evidence for speciation with gene flow in Heliconius butterflies. Genome Research, 23, 1817–1828.
Mavarez, J., Salazar, C. A., Bermingham, E., Salcedo, C., Jiggins, C. D. y Linares, M. (2006). Speciation by hibridization in Heliconius butterflies. Nature, 441(15), 868-871.
Mayr, E. (1942). Systematics and the Origin of Species. Columbia University Press, New York. Merchan, A., Jiggins, C. y Linares, M. (2005). A narrow Heliconius cydno (Nymphalidae; Heliconiini) hybrid zone with differences in morph sex ratios. Biotropica, 37 (1), 119-128.
Muñoz-Ortiz, A. (2004). Evidencia de la especiación incipiente entre Heliconius erato venus y Heliconius erato chestertonii. Tesis de Maestría. Departamento de Ciencias Biológicas. Facultad de Ciencias, Universidad de Los Andes, Bogotá, 81 p.
Naisbit, R.E., Jiggins, C.D., Linares, M. y Mallet, J. (2002). Hybrid sterility, Haldane's rule, and speciation in Heliconius cydno and H. melpomene. Genetics, 161, 1517-1526.
Neukirchen, W. (1991). Polymorphie und systematik von Heliconius longarena Hewitson (Lepidoptera: Nymphalidae). Entomologische Zeitschrift, 101 (3), 33-52.
Rosser, N., Kozak, K.M., Phillimore , A.B. y Mallet, J. (2015). Extensive range overlap between heliconiine sister species: evidence for sympatric speciation in butterflies? BMC Evolutionary Biology, 15, 125.
Rosser, N., Freitas, A., Huertas, B., Joron, M., Lamas, G., Merot, C., Simpson, F., Willmott, K.R., Mallet, J. y Dasmahapatra, K. (2019). Cryptic speciation associated with geographic and ecological divergence in two Amazonian Heliconius butterflies. Zoological Journal of the Linnean Society, 186, 233-249.
Salazar, C.A., Jiggins, C.D., Arias, C.F., Tobler, A., Bermingham, E. y Linares, M. (2005). Hybrid incompatibility is consistent with a hybrid origin of Heliconius heurippa Hewitson from its close relatives Heliconius cydno Doubleday and Heliconius melpomene Linnaeus. J Evol Biol., 18, 247-256.
Shapiro, B.J., Leducq, J.B. y Mallet, J. (2016). What is speciation? Plos Genetics, 12 (3), e1005860.
Sheppard, P.M., Turner, J.R.G., Brown, K.S., Benson, W.W. y Singer, M.C. (1985). Genetics and the evolution of Mullerian mimicry in Heliconius butterflies. Phil. Trans. R. Lond. B., 308, 433-610.
Stebbins, G.L. (1958). The inviability, weakness, and sterility of interspecific hybrids. Advances in Genetics, 9, 147-215.
The Heliconius Genome Consortium. (2012). Butterfly genome reveals promiscuous exchange of mimicry adaptations among species. Nature, 487 (7405), 94-98.
Turner, J.R.G. (1977). Butterfly mimicry: the genetical evolution of an adaptation. Evolutionary biology 10, 163-206.
Wahlberg, N., Braby, M., Brower, A., Lee, R., Nylin, S., Pierce, N., Sperling, F., Vila, R., Warren, A. y Zakharov, E. (2005). Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proc Roy Soc Lond., 272, 1577-1586.
Willmott, K.R., Constantino, L.M., y Hall, J.P.W. (2001). A review of Colobura (Lepidoptera: Nymphalidae) with comments on larval and adult ecology and description of a sibling species. Annals of the Entomological Society of America, 94 (2), 185-196.
Arias, C.F., Muñoz, A.G., Jiggins, C.D., Mavarez, J., Bermingham, E. y Linares, M. (2008) A bimodal hybrid zone provides evidence for incipient speciation in Heliconius butterflies. Molecular Ecology, 17, 4699-4712.
Audesirk, T. y Audesirk, G. (1996). El origen de las especies. Pp. 46-58. En: Audesirk, T. & Audesirk, G. (Ed). Evolución y ecología. Cuarta Edición. Prentice Hall Hispanoamericana S.A. México. 407 p.
Barton, N.H. y Hewitt, G.M. (1989). Adaptation, speciation and hybrid zones. Nature, 341, 497-503.
Brower, A.V. (1996). A new mimetic species of Heliconius (Lepidoptera: Nymphalidae), from southeastern Colombia revealed by cladistic analysis of mithochondrial DNA sequences. Zoological Journal of the Linnean Society, 116, 317-332.
Brower, A.V. (2011). Hybrid speciation in Heliconius butterflies? A review and critique of the evidence. Genetica, 139, 589-609.
Brown, K.S. y Benson, W. (1975). West Colombian biogeography. Notes on Heliconius hecalesia and H. sapho (Nymphalidae). Journal of the lepidopterists´ Society, 29 (4), 199-212.
Brown, K.S. (1975). Geographical patterns of evolution in Neotropical Lepidoptera. Systematics and derivation of known and new Heliconiini (Nymphalidae: Nymphalinae). Journal of Entomology
(B), 44 (3), 201-242.
Brown, K.S. (1981). The biology of Heliconius and related genera. Ann. Rev. Entomol., 26, 427-456.
Brown, K.S., Emmel, T.C., Eliazar, P.J. y Suomalainen, E. (1992). Evolutionary patterns in chromosome numbers in Neotropical Lepidoptera. I. Chromosomes of the Heliconiini. Hereditas, 117, 109–125.
Constantino, L.M. y Salazar, J. (1998). Natural hybridization of Heliconius cydno Doubleday from Western Colombia (Lepidoptera: Nymphalidae: Heliconiinae). Bol. Cient. Mus. Hist. Nat. U. Caldas, 2, 41-45.
Constantino, L.M., Gil, Z.N. y Corredor, G. (2005). Estudio del polimorfismo cromático en Laparus doris obscurus y determinación de la frecuencia fenotípica mediante cruzamientos intraespecíficos de tres poblaciones en el departamento del Valle y Caldas, Colombia (Nymphalidae: Heliconinae). Bol. Cient. Mus. Hist. Nat Universidad de Caldas,9, 222-237.
Constantino, L.M. y Salazar, J.A. (2010). A review of the Philaethria dido species complex (Lepidoptera. Nymphalidae: Heliconiinae) and description of three new sibling species from Colombia and Venezuela. Zootaxa, 2720, 1- 27.
Constantino, L.M. (2006). Redescubrimiento de Heliconius weymeri submarginatus, 98 años después de su aparente extinción y estudio de su complejo mimético con Heliconius hecubacrispus en el occidente de Colombia. Bol. Cient. Mus. Hist. Nat. Universidad de Caldas, 10, 138-197.
Constantino, L.M. (2016). Análisis morfológicos, moleculares y biogeográficos en la validación de nuevas especies y resolución de problemas taxonómicos en Lepidoptera. Memorias 43 Congreso Sociedad Colombiana de Entomología, Socolen. Manizales, 43, 209-228.
Dasmahapatra, K.K., Elias, M., Hill, R.I., Hoffman, J. y Mallet, J. (2010). Mitochondrial DNA barcoding detects some species that are real, and some that are not. Molecular Ecology Resources, 10, 264-273.
Dobzhansky, T. (1937). Genetics and the origin of species. Columbia University Press, New York.
Ebach, M.C. y Holdrege, C. (2005). DNA barcoding is no substitute for taxonomy. Nature, 434, 697-697.
Edelman, N.B., Frandsen, P.B., Miyagi, M., Clavijo, B., Davey, J., Dikow, R.B., García-Accinelli, G., Van Belleghem, S.M., Patterson, N., Neafsey, D. E., Challis, R., Kumar, S., Moreira, G.R.P., Salazar, C., Chouteau, M., Counterman, B.A., Papa, R., Blaxter, M., Reed, R.D., ... Mallet, J. (2019). Genomic architecture and introgression shape a butterfly radiation. Science, 366 (6465), 594-599.
Elias, M., Hill, R.I., Willmott, K.R., Dasmahapatra, K.K., Brower, A. V., Mallet, J. y Jiggins, C.D. (2007). Limited performance of DNA barcoding in a diverse community of tropical butterflies. Proceedings of the Royal Society of London B: Biological Sciences, 274 (1627), 2881-2889.
Gilbert, L.E. (2003). Adaptive novelty through introgression in Heliconius wing patterns: evidence for a shared genetic "toolbox" from synthetic hybrid zones and a theory of diversification. Ecology and Evolution Taking Flight: Butterflies as Model Systems. Edited by: Boggs C.L., Chicago: University of Chicago Press, 281-318.
Giraldo, C.E. (2015). La taxonomía integrativa en la resolución de problemas taxonómicos complejos en insectos: un caso de estudio en mariposas diurnas neotropicales. Libro de Memorias, 42 Congreso Sociedad Colombiana de Entomología, Medellín.,42, 142- 162.
Hendry, A.P. (2009). Speciation. Nature, 458, 162-164.
Jiggins, C.D., McMillan, W.O., Neukirchen, W. y Mallet, J. (1996). What can hybrid zones tell us about speciation? The case of Heliconius erato and H. himera (Lepidoptera: Nymphalidae). Biol J Linn Soc., 59, 221-242.
Jiggins, C.D. y McMillan, W.O. (1997). The genetics basis of an adaptive radiation: warning colour in two Heliconius species. Proc. R. Soc. Lond. B., 264, 1167-1175.
Jiggins, C.D. y Mallet, J. (2000). Bimodal hybrid zones and speciation. Trends Ecol Evol., , 15, 250-255.
Jiggins, C.D., Naisbit, R.E., Coe, R.L. y Mallet, J. (2001). Reproductive isolation caused by colour pattern mimicry. Nature, 411, 302-305.
Jiggins, C.D., Salazar, C., Linares, M. y Mavarez, J. (2008). Hybrid trait speciation and Heliconius butterflies. Speciation special issue of Phil. Trans. Royal. Soc., 363, 3047-3054.
Kozak, K.M., Wahlberg, N., Neild, A.F., Dasmahapatra, K.K., Mallet, J. y Jiggins, C.D. (2015). Multilocus species trees show the recent adaptive radiation of the mimetic Heliconius butterflies. Systematic Biology 64, 505-524.
Kronforst., M.R., Young, L.G., Blume, L. M. y Gilbert, L.E. (2006). Multilocus analysis of admixture and introgression among hybridizing Heliconius butterflies. Evolution, 60, 1254-1268.
Lamas, G. (2004). Heliconiinae pp. 261-274. In. Lamas, G. (Ed.), Atlas of Neotropical Lepidoptera, Check list: Part 4A, HesperoideaPapilionoidea. Association for Tropical Lepidoptera, Gainesville. 439 pp.
Linares, M. (1996). The genetics of the mimetic coloration in the butterfly Heliconius cydno weymeri. Heredity, 87, 142-149.
Mallet, J. y Barton, N. H. (1989). Strong natural selection in a warning colour hybrid zone. Evolution, 43, 421-431.
Mallet, J. (1993). Speciation, radiation, and color pattern evolution in Heliconius butterflies: evidence from hybrid zones. Hybrid Zones and the Evolutionary Process. Edited by: Harrison RG. New York: Oxford University Press, 226-260.
Mallet, J. (1995). A species definition for the Modern Synthesis. Tree, 10, 294-298.
Mallet, J. y Gilbert, L.E., 1995.- Why are there so many mimicry rings? Correlations between habitat, behaviour and mimicry in Heliconius butterflies. Biol J Linn Soc., 55, 159-180.
Mallet, J. (1999). Causes and consequences of a lack of coevolution in Müllerian mimicry. Evolutionary Ecology, 13, 777-806.
Mallet, J. (2007). Hybrid speciation. Reviews. Nature, 446, 279-283.
Mallet, J. (2013). Species, concept of. Encyclopedia of Biodiversity. Volume 6, 679-691.
Mallet, J. y Willmott, K. (2003). Taxonomy: renaissance or Tower of Babel? Trends Ecol. Evol., 18, 57-59.
Mallet, J., Beltran, M., Neukirchen, W. y Linares, M. (2007). Natural hybridization in Heliconiine butterflies: the species boundary as a continuum. BMC Evolutionary Biology. 7,1-16.
Martin, S.H., Dasmahapatra, K.K., Nadeau, N.J., Salazar, C., Walters, J.R., Simpson, F., Blaxter, M., Manica, A., Mallet, J. y Jiggins, C.D. (2013). Genome-wide evidence for speciation with gene flow in Heliconius butterflies. Genome Research, 23, 1817–1828.
Mavarez, J., Salazar, C. A., Bermingham, E., Salcedo, C., Jiggins, C. D. y Linares, M. (2006). Speciation by hibridization in Heliconius butterflies. Nature, 441(15), 868-871.
Mayr, E. (1942). Systematics and the Origin of Species. Columbia University Press, New York. Merchan, A., Jiggins, C. y Linares, M. (2005). A narrow Heliconius cydno (Nymphalidae; Heliconiini) hybrid zone with differences in morph sex ratios. Biotropica, 37 (1), 119-128.
Muñoz-Ortiz, A. (2004). Evidencia de la especiación incipiente entre Heliconius erato venus y Heliconius erato chestertonii. Tesis de Maestría. Departamento de Ciencias Biológicas. Facultad de Ciencias, Universidad de Los Andes, Bogotá, 81 p.
Naisbit, R.E., Jiggins, C.D., Linares, M. y Mallet, J. (2002). Hybrid sterility, Haldane's rule, and speciation in Heliconius cydno and H. melpomene. Genetics, 161, 1517-1526.
Neukirchen, W. (1991). Polymorphie und systematik von Heliconius longarena Hewitson (Lepidoptera: Nymphalidae). Entomologische Zeitschrift, 101 (3), 33-52.
Rosser, N., Kozak, K.M., Phillimore , A.B. y Mallet, J. (2015). Extensive range overlap between heliconiine sister species: evidence for sympatric speciation in butterflies? BMC Evolutionary Biology, 15, 125.
Rosser, N., Freitas, A., Huertas, B., Joron, M., Lamas, G., Merot, C., Simpson, F., Willmott, K.R., Mallet, J. y Dasmahapatra, K. (2019). Cryptic speciation associated with geographic and ecological divergence in two Amazonian Heliconius butterflies. Zoological Journal of the Linnean Society, 186, 233-249.
Salazar, C.A., Jiggins, C.D., Arias, C.F., Tobler, A., Bermingham, E. y Linares, M. (2005). Hybrid incompatibility is consistent with a hybrid origin of Heliconius heurippa Hewitson from its close relatives Heliconius cydno Doubleday and Heliconius melpomene Linnaeus. J Evol Biol., 18, 247-256.
Shapiro, B.J., Leducq, J.B. y Mallet, J. (2016). What is speciation? Plos Genetics, 12 (3), e1005860.
Sheppard, P.M., Turner, J.R.G., Brown, K.S., Benson, W.W. y Singer, M.C. (1985). Genetics and the evolution of Mullerian mimicry in Heliconius butterflies. Phil. Trans. R. Lond. B., 308, 433-610.
Stebbins, G.L. (1958). The inviability, weakness, and sterility of interspecific hybrids. Advances in Genetics, 9, 147-215.
The Heliconius Genome Consortium. (2012). Butterfly genome reveals promiscuous exchange of mimicry adaptations among species. Nature, 487 (7405), 94-98.
Turner, J.R.G. (1977). Butterfly mimicry: the genetical evolution of an adaptation. Evolutionary biology 10, 163-206.
Wahlberg, N., Braby, M., Brower, A., Lee, R., Nylin, S., Pierce, N., Sperling, F., Vila, R., Warren, A. y Zakharov, E. (2005). Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proc Roy Soc Lond., 272, 1577-1586.
Willmott, K.R., Constantino, L.M., y Hall, J.P.W. (2001). A review of Colobura (Lepidoptera: Nymphalidae) with comments on larval and adult ecology and description of a sibling species. Annals of the Entomological Society of America, 94 (2), 185-196.
