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Marquez-P, J. ., & Martinez-H, N. . (2020). Population structure of Morpho helenor peleides Kollar, 1850 (Lepidoptera: Nymphalidae) in a tropical dry forest landscape, department of Bolivar, Colombia. Boletín Científico. Centro De Museos, 24(2), 169–189. https://doi.org/10.17151/bccm.2020.24.2.12
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Authors
Johana Marquez-P
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Neis Martinez-H
Abstract
Considering the high destruction rate of forest in the Caribbean of Colombia and the lacking of available information about population ecology of butterflies from the genus Morpho, we analyzed the variation of the population structure of Morpho helenor peleides Kollar, 1850 in a Tropical Dry Forest (TDF) landscape on “Montes de María”, in the Caribbean of Colombia. Methodology. For the butterfly study, we used the capture-mark-recapture method (CMR) on seven stations, in which we installed Van Someren-Rydon traps, baited with a mixture of fermented fruits. Results. The total abundance was 287 individuals, with 59 recaptures (20,5%). The population size we obtained was between 8 and 339 individuals. Sexual proportion male-female was 3:1. We captured a higher percentage of new butterflies (41,8%) and intermediate butterflies (38,6%). The maximum residence time we registered was 38 days. We calculated total distances of flight between 149 and 1,231; although females presented the highest average distances (303.7±189.4m), there was not statistical significance in the distances of flight between sexes (U= 265,5, p>0,05). Conclusions. The population structure of M. helenor peleides presented a spatio-temporal variation in the dry forest landscape we studied. Additionally, we determined that natural canopy cover plays an important role in the temporal dynamic and distribution of the abundance of this butterfly group in tropical dry forest, and it is emphasized the importance of this ecosystem in the conservation of this species.
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References
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Andrade-C, M.G., Campos, S.L., Gonzales, M.I., Pulido, H.W. (2007). Santa María, Mariposas, Alas y Color: 100. ICN, Universidad Nacional de Colombia, Bogotá.
Basset, Y., Eastwood, R., Sam, L., Lohman, V., Treuer, T., Miller, S., Osorio, M. (2012). Cross-continental comparisons of butterfly assemblages in tropical rainforests: implications for biological monitoring. Insect Conservation and Diversity, 6:223–233.
Besold, J., Huck, S., Schmitt, T. (2008). Allozyme polymorphisms in the small heath, Coenonympha pamphilus: recent ecological selection or biogeographical signal?. Annales Zoologici
Fennici, 45: 217-228.
Brown, K. & Freitas, A.V. (2000). Atlantic Forest Butterflies: Indicators for Landscape conservation. Biotropica, 32(4b): 934-956.
Brussard, P.F., Ehrlich, P.R., Singer M.C. (1974). Adult movements and population structure in Euphydrias editha. Evolution, 28: 408-415.
Campos-Salazar, L.R., Gómez, B.J., Andrade-C, M.G. (2011). Mariposas (Lepidoptera: Hesperioidea-Papilionoidea) de las áreas circundantes a las Ciénagas del Departamento de Córdoba,
Colombia. Revista Acad. Colomb. Ci. Exact., 35(134):45-60.
Castaño Uribe, C.S. (1999). Serranías de Colombia. Banco de Occidente I/M Editores. Cali, 329.
Chazot, N., Panaras, S., Zilbermann, N., Blandin, P., Le Poul, Y., Cornette, R., Elias, M., Debat, V. (2016). Morpho morphometrics: Shared encestry and selection drive the evolution of wings size and shape in Morpho butterflies. Evolution, 70(1):181-194.
Clark, L.R., Geier, P.W., Hugues, R.D., Morris, R.F. (1978). The Ecology of insect Populations in Theory and Practice, 4th ed.; The English Language Book Society and Methuen Book Society: London, UK, pp 1-24, ISBN 416766900.
Collinge, S.K., Prudic, K.L., Oliver, J.C. (2003). Effects of local habitat characteristics and landscape context on grassland butterfly diversity. Conservation Biology, 17(1):178-187.
Constantino, L.M. & Corredor, G. (2004). The biology and morphology of the early stages of Morpho macrophtalmus and M. peleides telamon (Nymphalidae: Morphinae) from western Colombia. Bol. Cient. Mus. Hist. Nat. U. de Caldas, 8:201-209.
Devries, P.J., Murray, D., Lande, R. (1997). Species diversity in vertical, horizontal, and temporal dimensions of a fruit-feeding butterfly community in an Ecuadorian rainforest. Biol. J. Linn. Soc., 62: 343-364.
Devries, P.J. (1987). The Butterflies of Costa Rica and their Natural History, Papilionidae, Pieridae, Nymphalidae. Princeton Univ Press, 327 p.
Díaz, A.C., Barbosa, C.E., De la Ossa, J. (1986). Aspectos ecológicos y etológicos de primates con énfasis en Alouatta Seniculus (Cebidae), de la región de Coloso, Serranía de San Jacinto (Sucre), Costa Norte de Colombia. Caldasia, 14:709-741.
Di Rienzo, J. A., Casanoves, F., Balzarini, M. G., Gonzalez, L., Tablada, M., Robledo C.W. (2019). InfoStat (versión 2019). Centro de Transferencia InfoStat, FCA, Universidad Nacional de Córdoba, Argentina. Recuperado de http://www.infostat.com.ar.
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Fagua, G. & Ruiz, N. (1996). Relaciones de herbivoría entre papiliónidas (Lepidoptera) y especies de Aristolochia (Aristolochiaceae). En: Andrade, C.G., García, A.G., Fernández, F. (eds.). Insectos de Colombia. Centro Editorial Javeriano (ceja). Revista Acad. Colomb. Ci. Exact., 541 p.
Freire, G., Nascimento, A.R., Konstantino, I.M., Diniz, I.R. (2014). Temporal ocurrence of two Morpho butterflies (Lepidoptera: Nymphalidae): influence of weather and food resources. Environmental Entomology, 43:274-282.
Freitas, A. & Ramos, R. (2001). Population biology of Parides Anchises nephalion (Papilionidae) in a coastal site in southeast Brazil. Brazi. J. Biol., 61(4):623-630.
Garcia, C.R., Gallusser, S., Lachaume, G., Blandin, P. (2014). The ecology and life cycle of the Amazonian Morho cisseis phanodemus Hewitson, 1869, whit a comparative review of early stages in the genus Morpho (Lepidoptera: Nymphalidae: Morphinae). Tropical Lipidoptera Research, 24(2):67-80.
García, R.C., Constantino, L.M., Heredia, M.D., Kattan, G. (2002). Guía de campo: Mariposas comunes de la cordillera central de Colombia. p.130, ISBN:958-33-3190-2.
Hammer. (2013). PAST Paleontological Statistics Version 3.0: Reference Manual University of Oslo.
Heyer, W.R., Donelly, M.A., Mcdiarmid, R.W., Hayec, L.A, Foster, M.S. (1994). Measuring and monitoring biological diversity: standard methods for amphibians. Smithsonian institution. Washington, 364 p.
Jolly, G.M. (1965). Explicit estimates from capture-recapture data with both death and immigration stochastic model. Biometrika, 1: 225-247.
Krebs, C.J., Kenney, A.J. (1998). Ecology Methodology. Department of zoology, University of British Columbia, Vancouver, B.C Canada V6T 1Z4.
Kremen, C. (1994). Biological inventory using target taxa a case study of the butterflies of Madagascar. Ecological Applications, 4(3): 407-422.
Lemos, J.A., Rojas, R.I., Zúñiga, J.J. (2005). Técnicas para el estudio de poblaciones de fauna silvestre. Jiménez editores. Universidad Nacional autónoma de México, México D.F, 157 p.
Martínez, N., García, S., Gutierrez, M., Sanjuán, S., Contreras, C. (2010). Composición y estructura de la fauna de escarabajos (Insecta: Coleoptera) atraídos por trampas de luz en La Reserva Ecológica de Luriza, Atlántico, Colombia. Boletín Sociedad Entomológica Aragonesa, 373-381 p.
Molleman, F., Ding, J., Wang, J.L., Zwaan, B.J., Carey, J.R., Brakefield, P.M. (2008). Adult diet affects lifespan and reproduction of the fruit-feeding butterfly Charaxes fulvescens. Entomologia Experimentalis et Applicata, 129(1): 54–65.
Ochoa, S.J.A., Escorcia, D.K., Duran, J., Prieto, C. (2018). Population Structure of Morpho rhodopteron, Godman & Salvin (Lepidoptera: Nymphalidae) an Endemic Butterfly from the Sierra Nevada De Santa Marta, Colombia. J. Lepid. Soc. 72(4): 320-326.
Palanca, A. (1975). Lepidópteros y tiempo atmosférico. Graellsia, 31:61-77.
Pizano, C., García, H. (2014). El bosque seco tropical en Colombia. Instituto de Investigación de Recursos Biológicos.
Pollard, E. (1988). Temperature, rainfall and butterfly numbers. J. Appl. Eco., 819-828.
Prieto, C., Takegami, C., Rivera, J.M. (2005). Estructura poblacional de Morpho sulkowskyi Kollar, 1850 (Lepidoptera: Nymphalidae) en un sector de la cordillera occidental, departamento del
Cauca (Colombia). Entomotropica, 20(1):15-22.
Ramos, R., Freitas, A. (1999). Population biology, wing color variation and ecological plasticity in Heliconius erato phyllis (Nymphalidae). J. Lepid. Soc., 53:11-21.
Rangel, J., Carvajal, C.J. (2012). Clima de la región Caribe colombiana. Colombia Diversa, Biótica XII. Región Caribe Colombia. Bogotá, Instituto de Ciencias Naturales Universidad Nacional de Colombia: 67–129.
Ribeiro, D.B., Freitas, A.V.L. (2011). Large-sized insects show stronger seasonality tan small-sized ones: a case study of fruit-feeding butterflies. Biol. J. Linn. Soc., 104:820-827.
Ribeiro, D.B., Prado, P.I., Brown J.R., K.S., Freitas, A.V.L. (2008). Additive partitioning of butterfly diversity in a fragmented landscape importance of scale and implications for conservation. Diversity and Distributions, 14: 961-968.
Rodríguez, J.V., Rueda, J.V., Gutiérrez, T.D. (2008). Guía ilustrada de la fauna del Santuario de Vida Silvestre Los Besotes, Valledupar, Cesar, Colombia. Serie de guías tropicales de campo
Nº7, Conservación Internacional. Editorial Panamericana. Bogotá, Colombia.
Ruiz, E., Vásquez, J., Zárate, R., Pinedo, J. (2015). Aspectos biológicos de Morpho helenor theodorus (Fruhstorfer) (Lepidoptera: Nymphalidae: Morphinae) y Mechanitis polymnia (Linnaeus) (Lepidoptera: Nymphalidae: Ithominae), en la Amazonia baja del Perú. Instituto de Investigaciones de la Amazonia Peruana (IIAP), 24(1):45-54.
Sampedro, A., Álvarez, A., Domínguez, L.M., Herrera, I. (2013). Especies promisorias para el ecoturismo en “Campo Aventura Roca Madre”, Toluviejo-Sucre, Colombia. Revista MVZ Córdoba, 18(1):3387-3398.
Schaeffer, P.V., Oliveira, M.N., Piccoli, R.H. (2019). Population structure and natural history of the South American fruit-feeding butterfly Taygetis ypthima (Lepidoptera:Nymphalidae). Austral Entomology, 58(4): 753-761.
Seber, G.A. (1965). A note on the multiple-recapture census. Biometrika, 52: 249-259.
Strickler, G. (1959). Use of the densitometer to estimate density of forest canopy on permanent sample plots. USDA For Serv Research.
Uehara-Prado, M., Brown J.R., K.S., Freitas, A.V.L. (2007). Species richness, compositionand abundance of fruit-feeding butterflies in the Brazilian Atlantic Forest: comparison between
afragmented and a continuous landscape. Glob. Ecol. Biogeogr., 16: 43-54.
Vargas, M., Martínez, N., Gutiérrez, L.C., Prince, S. (2011). Riqueza y abundancia de Hesperioidea y Papilionoidea (Lepidóptera) en la Reserva Natural Las Delicias, Santa Marta, Magdalena, Colombia. Acta Biol. Colomb., 16(1):43-60.
Vargas, M., Prince, S., Martínez, N. (2012). Estructura poblacional de Heliconius erato hydara Hewitson, 1867 (Lepidoptera: Nymphalidae) en la Reserva Campesina la Montaña, departamento del Atlántico, Colombia. Boletín Sociedad Entomológica Aragonesa, 51:273-281.
Vu, L.V. (2009). Diversity and similarity of butterfly communities in five different habitat types at Tamdao National Park Vietnam. Journal of Zoology, 227: 15-22.
Wilson, K., Hardy, I.C.W. (2002). Statistical analysis of sex ratios: An introduction. In Sex Ratios: Concepts and Research Methods, 1st ed, Hardy ICW, ed, Cambridge University Press: New
York, NY, USA, pp. 48-92, ISBN 978052181896.
Wolda, H. (1978). Seasonal fluctuation in rainfall, food and abundance of tropical insects. J. Anim. Ecol., 47:369-381.
Young, A.M., Muyshondt, A. (1973). The biology of Morpho peleides in Central America. Caribb. J. Sci., 13:1- 49.
Young, A.M., Thomason, J.H. (1974). The demography of a confined population of the butterfly Morpho peleides during a tropical dry season. Stud. Neotrop. Fauna Environ., 9:1-34.
Young, A.M. (1982). Notes on the natural history of Morpho granadensis polybaptus Butler (Nymphalidae: Morphinae), and its relation to that of Morpho peleides limpida Butler. J.N.Y Entomol. Soc., 90:35-54.
Ancona, S., Dénes, F.V., Krüger, O., Székely, T., Beissinger S.R. (2017). Estimating adult sex ratios in nature. Proceedings of the Royal Society B: Biological Sciences, 372(1729), [20160313].
Andrade-C, M.G., Campos, S.L., Gonzales, M.I., Pulido, H.W. (2007). Santa María, Mariposas, Alas y Color: 100. ICN, Universidad Nacional de Colombia, Bogotá.
Basset, Y., Eastwood, R., Sam, L., Lohman, V., Treuer, T., Miller, S., Osorio, M. (2012). Cross-continental comparisons of butterfly assemblages in tropical rainforests: implications for biological monitoring. Insect Conservation and Diversity, 6:223–233.
Besold, J., Huck, S., Schmitt, T. (2008). Allozyme polymorphisms in the small heath, Coenonympha pamphilus: recent ecological selection or biogeographical signal?. Annales Zoologici
Fennici, 45: 217-228.
Brown, K. & Freitas, A.V. (2000). Atlantic Forest Butterflies: Indicators for Landscape conservation. Biotropica, 32(4b): 934-956.
Brussard, P.F., Ehrlich, P.R., Singer M.C. (1974). Adult movements and population structure in Euphydrias editha. Evolution, 28: 408-415.
Campos-Salazar, L.R., Gómez, B.J., Andrade-C, M.G. (2011). Mariposas (Lepidoptera: Hesperioidea-Papilionoidea) de las áreas circundantes a las Ciénagas del Departamento de Córdoba,
Colombia. Revista Acad. Colomb. Ci. Exact., 35(134):45-60.
Castaño Uribe, C.S. (1999). Serranías de Colombia. Banco de Occidente I/M Editores. Cali, 329.
Chazot, N., Panaras, S., Zilbermann, N., Blandin, P., Le Poul, Y., Cornette, R., Elias, M., Debat, V. (2016). Morpho morphometrics: Shared encestry and selection drive the evolution of wings size and shape in Morpho butterflies. Evolution, 70(1):181-194.
Clark, L.R., Geier, P.W., Hugues, R.D., Morris, R.F. (1978). The Ecology of insect Populations in Theory and Practice, 4th ed.; The English Language Book Society and Methuen Book Society: London, UK, pp 1-24, ISBN 416766900.
Collinge, S.K., Prudic, K.L., Oliver, J.C. (2003). Effects of local habitat characteristics and landscape context on grassland butterfly diversity. Conservation Biology, 17(1):178-187.
Constantino, L.M. & Corredor, G. (2004). The biology and morphology of the early stages of Morpho macrophtalmus and M. peleides telamon (Nymphalidae: Morphinae) from western Colombia. Bol. Cient. Mus. Hist. Nat. U. de Caldas, 8:201-209.
Devries, P.J., Murray, D., Lande, R. (1997). Species diversity in vertical, horizontal, and temporal dimensions of a fruit-feeding butterfly community in an Ecuadorian rainforest. Biol. J. Linn. Soc., 62: 343-364.
Devries, P.J. (1987). The Butterflies of Costa Rica and their Natural History, Papilionidae, Pieridae, Nymphalidae. Princeton Univ Press, 327 p.
Díaz, A.C., Barbosa, C.E., De la Ossa, J. (1986). Aspectos ecológicos y etológicos de primates con énfasis en Alouatta Seniculus (Cebidae), de la región de Coloso, Serranía de San Jacinto (Sucre), Costa Norte de Colombia. Caldasia, 14:709-741.
Di Rienzo, J. A., Casanoves, F., Balzarini, M. G., Gonzalez, L., Tablada, M., Robledo C.W. (2019). InfoStat (versión 2019). Centro de Transferencia InfoStat, FCA, Universidad Nacional de Córdoba, Argentina. Recuperado de http://www.infostat.com.ar.
Ehrlich, P.R. & Gilbert, L.E. (1973). Population structure and dynamics of the tropical butterfly Heliconius ethilla. Biotropica, 5, 69-82.
Fagua, G. & Ruiz, N. (1996). Relaciones de herbivoría entre papiliónidas (Lepidoptera) y especies de Aristolochia (Aristolochiaceae). En: Andrade, C.G., García, A.G., Fernández, F. (eds.). Insectos de Colombia. Centro Editorial Javeriano (ceja). Revista Acad. Colomb. Ci. Exact., 541 p.
Freire, G., Nascimento, A.R., Konstantino, I.M., Diniz, I.R. (2014). Temporal ocurrence of two Morpho butterflies (Lepidoptera: Nymphalidae): influence of weather and food resources. Environmental Entomology, 43:274-282.
Freitas, A. & Ramos, R. (2001). Population biology of Parides Anchises nephalion (Papilionidae) in a coastal site in southeast Brazil. Brazi. J. Biol., 61(4):623-630.
Garcia, C.R., Gallusser, S., Lachaume, G., Blandin, P. (2014). The ecology and life cycle of the Amazonian Morho cisseis phanodemus Hewitson, 1869, whit a comparative review of early stages in the genus Morpho (Lepidoptera: Nymphalidae: Morphinae). Tropical Lipidoptera Research, 24(2):67-80.
García, R.C., Constantino, L.M., Heredia, M.D., Kattan, G. (2002). Guía de campo: Mariposas comunes de la cordillera central de Colombia. p.130, ISBN:958-33-3190-2.
Hammer. (2013). PAST Paleontological Statistics Version 3.0: Reference Manual University of Oslo.
Heyer, W.R., Donelly, M.A., Mcdiarmid, R.W., Hayec, L.A, Foster, M.S. (1994). Measuring and monitoring biological diversity: standard methods for amphibians. Smithsonian institution. Washington, 364 p.
Jolly, G.M. (1965). Explicit estimates from capture-recapture data with both death and immigration stochastic model. Biometrika, 1: 225-247.
Krebs, C.J., Kenney, A.J. (1998). Ecology Methodology. Department of zoology, University of British Columbia, Vancouver, B.C Canada V6T 1Z4.
Kremen, C. (1994). Biological inventory using target taxa a case study of the butterflies of Madagascar. Ecological Applications, 4(3): 407-422.
Lemos, J.A., Rojas, R.I., Zúñiga, J.J. (2005). Técnicas para el estudio de poblaciones de fauna silvestre. Jiménez editores. Universidad Nacional autónoma de México, México D.F, 157 p.
Martínez, N., García, S., Gutierrez, M., Sanjuán, S., Contreras, C. (2010). Composición y estructura de la fauna de escarabajos (Insecta: Coleoptera) atraídos por trampas de luz en La Reserva Ecológica de Luriza, Atlántico, Colombia. Boletín Sociedad Entomológica Aragonesa, 373-381 p.
Molleman, F., Ding, J., Wang, J.L., Zwaan, B.J., Carey, J.R., Brakefield, P.M. (2008). Adult diet affects lifespan and reproduction of the fruit-feeding butterfly Charaxes fulvescens. Entomologia Experimentalis et Applicata, 129(1): 54–65.
Ochoa, S.J.A., Escorcia, D.K., Duran, J., Prieto, C. (2018). Population Structure of Morpho rhodopteron, Godman & Salvin (Lepidoptera: Nymphalidae) an Endemic Butterfly from the Sierra Nevada De Santa Marta, Colombia. J. Lepid. Soc. 72(4): 320-326.
Palanca, A. (1975). Lepidópteros y tiempo atmosférico. Graellsia, 31:61-77.
Pizano, C., García, H. (2014). El bosque seco tropical en Colombia. Instituto de Investigación de Recursos Biológicos.
Pollard, E. (1988). Temperature, rainfall and butterfly numbers. J. Appl. Eco., 819-828.
Prieto, C., Takegami, C., Rivera, J.M. (2005). Estructura poblacional de Morpho sulkowskyi Kollar, 1850 (Lepidoptera: Nymphalidae) en un sector de la cordillera occidental, departamento del
Cauca (Colombia). Entomotropica, 20(1):15-22.
Ramos, R., Freitas, A. (1999). Population biology, wing color variation and ecological plasticity in Heliconius erato phyllis (Nymphalidae). J. Lepid. Soc., 53:11-21.
Rangel, J., Carvajal, C.J. (2012). Clima de la región Caribe colombiana. Colombia Diversa, Biótica XII. Región Caribe Colombia. Bogotá, Instituto de Ciencias Naturales Universidad Nacional de Colombia: 67–129.
Ribeiro, D.B., Freitas, A.V.L. (2011). Large-sized insects show stronger seasonality tan small-sized ones: a case study of fruit-feeding butterflies. Biol. J. Linn. Soc., 104:820-827.
Ribeiro, D.B., Prado, P.I., Brown J.R., K.S., Freitas, A.V.L. (2008). Additive partitioning of butterfly diversity in a fragmented landscape importance of scale and implications for conservation. Diversity and Distributions, 14: 961-968.
Rodríguez, J.V., Rueda, J.V., Gutiérrez, T.D. (2008). Guía ilustrada de la fauna del Santuario de Vida Silvestre Los Besotes, Valledupar, Cesar, Colombia. Serie de guías tropicales de campo
Nº7, Conservación Internacional. Editorial Panamericana. Bogotá, Colombia.
Ruiz, E., Vásquez, J., Zárate, R., Pinedo, J. (2015). Aspectos biológicos de Morpho helenor theodorus (Fruhstorfer) (Lepidoptera: Nymphalidae: Morphinae) y Mechanitis polymnia (Linnaeus) (Lepidoptera: Nymphalidae: Ithominae), en la Amazonia baja del Perú. Instituto de Investigaciones de la Amazonia Peruana (IIAP), 24(1):45-54.
Sampedro, A., Álvarez, A., Domínguez, L.M., Herrera, I. (2013). Especies promisorias para el ecoturismo en “Campo Aventura Roca Madre”, Toluviejo-Sucre, Colombia. Revista MVZ Córdoba, 18(1):3387-3398.
Schaeffer, P.V., Oliveira, M.N., Piccoli, R.H. (2019). Population structure and natural history of the South American fruit-feeding butterfly Taygetis ypthima (Lepidoptera:Nymphalidae). Austral Entomology, 58(4): 753-761.
Seber, G.A. (1965). A note on the multiple-recapture census. Biometrika, 52: 249-259.
Strickler, G. (1959). Use of the densitometer to estimate density of forest canopy on permanent sample plots. USDA For Serv Research.
Uehara-Prado, M., Brown J.R., K.S., Freitas, A.V.L. (2007). Species richness, compositionand abundance of fruit-feeding butterflies in the Brazilian Atlantic Forest: comparison between
afragmented and a continuous landscape. Glob. Ecol. Biogeogr., 16: 43-54.
Vargas, M., Martínez, N., Gutiérrez, L.C., Prince, S. (2011). Riqueza y abundancia de Hesperioidea y Papilionoidea (Lepidóptera) en la Reserva Natural Las Delicias, Santa Marta, Magdalena, Colombia. Acta Biol. Colomb., 16(1):43-60.
Vargas, M., Prince, S., Martínez, N. (2012). Estructura poblacional de Heliconius erato hydara Hewitson, 1867 (Lepidoptera: Nymphalidae) en la Reserva Campesina la Montaña, departamento del Atlántico, Colombia. Boletín Sociedad Entomológica Aragonesa, 51:273-281.
Vu, L.V. (2009). Diversity and similarity of butterfly communities in five different habitat types at Tamdao National Park Vietnam. Journal of Zoology, 227: 15-22.
Wilson, K., Hardy, I.C.W. (2002). Statistical analysis of sex ratios: An introduction. In Sex Ratios: Concepts and Research Methods, 1st ed, Hardy ICW, ed, Cambridge University Press: New
York, NY, USA, pp. 48-92, ISBN 978052181896.
Wolda, H. (1978). Seasonal fluctuation in rainfall, food and abundance of tropical insects. J. Anim. Ecol., 47:369-381.
Young, A.M., Muyshondt, A. (1973). The biology of Morpho peleides in Central America. Caribb. J. Sci., 13:1- 49.
Young, A.M., Thomason, J.H. (1974). The demography of a confined population of the butterfly Morpho peleides during a tropical dry season. Stud. Neotrop. Fauna Environ., 9:1-34.
Young, A.M. (1982). Notes on the natural history of Morpho granadensis polybaptus Butler (Nymphalidae: Morphinae), and its relation to that of Morpho peleides limpida Butler. J.N.Y Entomol. Soc., 90:35-54.
