This is a list of scientific studies arranged by general topics and chronologically.



Diversity and Distribution

Systematics and Biogeography

Origin of Carica papaya

Wild form of Carica papaya

Sexual systems


Sex Chromosomes



Agronomy, Physiology, and Chemistry

Dated phylogenies (only studies including Caricaceae)

Crop domestication


For weblinks to the protologues use the search form.

Carvalho, F., Filer, D., Renner, S. S. 2014. Taxonomy in the electronic age and an e-monograph of the papaya family (Caricaceae) as an example. Cladistics, Early view. doi: : 10.1111/cla.12095 abstract.

McVaugh, R. 2001. Caricaceae. In: Flora Novo-Galiciana. A Descriptive Account of the Vascular Plants of Western Mexico, vol. 3 Ochnaceae to Losaceae. The University of Michigan Press, Ann Arbor, Michigan, pp. 71-73.

Badillo, V.M. 2001. Nota correctiva Vasconcellea St. Hill. y no Vasconcella (Caricaceae). Ernstia 10, 74-79.

Badillo, V.M. 2000. Carica L. vs. Vasconcella St.-Hil. (Caricaceae) con la rehabilitacion de este ultimo. Ernstia 10, 74-79.

Badillo, V.M. 1999. Nuevos registros para el Ecuador de especies del género Carica. Ernstia 9(3-4): 185-186.

Badillo, V.M. 1997. Neotipificación de Carica pubescens Lenné et Koch y de Carica quercifolia (St. Hil.) Hieron. y nuevos registros de la familia para Ecuador. Ernstia 6(4): 201-205.

Badillo, V.M. 1993. Caricaceae. Segundo Esquema. Rev. Fac. Agron. Univ. Cent. Venezuela 43, 1-111.

Diaz-Luna, C.L., Lomeli-Sención, J.A. 1992. Revisión del género Jarilla Rusby (Caricaceae). Acta Bot. Mexicana 20, 77-99. open access.

Badillo V.M., 1983. Caricaceae. In: Harling G. and Sparre B. (Eds.). Flora of Ecuador No. 20. Göteborg University, Göteborg, Sweden. 48 p. (buy)

Lorence, D.H., Colin, R.T. 1988. Carica cnidoscoloides (sp. nov.) and sect. Holostigma (sect. nov.) of Caricaceae from Southern Mexico. Syst. Bot. 13, 107-110. abstract

Badillo, V.M. 1983. Caricaceae. Segundo Esquema. Rev. Fac. Agron. Univ. Cent. Venezuela 43, 1-111.

Moreno, N.P. 1980. Caricaceae. Flora de Veracruz, Fasc. 10. Instituto de Ecología, Xalapa.

Badillo, V.M. 1974. Notas adicionales a la monografia de la familia Caricaceae. Revista Fac. Agron. (Maracay) 7(3): 22-23.

Badillo, V.M. 1971. Monografía de la família Caricaceae. Asociación de profesores, Universidad Central de Venezuela, Facultad de Agronomía, Maracay.

Badillo, V.M. 1967. Esquema de las Caricaceae. Revista Fac. Agron. (Maracay) 4(7): 245-272.

Badillo, V.M. 1967. Inventario anotado de las Caricaceae hasta hoy tenidas como validas. Revista Fac. Agron. (Maracay) 4(2): 3-21.

Badillo, V.M. 1967. Cinco nuevas especies ecuatorianas y colombianas del genero Carica. Revista Fac. Agron. (Maracay) 4(2): 67-91.

Badillo, V.M. 1966. Dos nuevas especies ecuatorianas del genero Carica. Revista Fac. Agron. (Maracay) 4(1): 57-64.


Tshimala-Tshibangu, N. & Ndjigba; J.D. 1998. Utilisations des produits forestiers autres que le bois (PFAB) au Cameroun. Cas du projet forestier du Mont Koupé. Tropicultura, 16-17 (2): 70-79. open access

Terashima, H. 1991. AFlora: Catalog of Useful Plants of Tropical Africa. African Study Monographs, Supp!. 16: 1-195

Diversity and Distribution

Scheldeman, X., Willemen, L., Coppens d'Eeckenbrugge, G., Romeijn-Peeters, E., Restrepo, M.T., Romero Motoche, J., Jiménez, D., Lobo, M., Medina, C.I., Reyes, C., Rodríguez, D., Ocampo, J.A., Damme, P., Goetgebeur, P. 2007. Distribution, diversity and environmental adaptation of highland papayas (Vasconcellea spp.) in tropical and subtropical America. Biodivers. Conserv. 16, 1867-1884. doi: 10.1007/978-1-4020-6444-9_19 abstract.

Rodríguez, D., Marín, C., Quecan, H., Ortiz, R., 2005. Áreas potenciales para colectas del género Vasconcellea Badillo en venezuela. Bioagro 17, 3-10. open access

Hofer, U., Bersier, L., Borcard, D. 2000. Ecotones and gradient as determinants of herpetofaunal community structure in the primary forest of Mount Kupe, Cameroon. Journal of Tropical Ecology 16: 517-533. open access

Systematics and Biogeography

Carvalho, F. and Renner, S.S. 2012. A dated phylogeny of the papaya family (Caricaceae) reveals the crop's closest relatives and the family's biogeographic history. Mol. Phylogenet. Evol. 65(1): 46-53. doi: 10.1016/j.ympev.2012.05.019 open access

Kyndt, T., Van Droogenbroeck, B., Romeijn-Peeters, E., Romero-Motochi, J.P., Scheldeman, X., Goetghebeur, P., Van Damme, P., Gheysen, G. 2005a. Molecular phylogeny and evolution of Caricaceae based on rDNA internal transcribed spacers and chloroplast sequence data. Mol. Phylogenet. Evol. 37(2): 442-459. doi: 10.1016/j.ympev.2005.06.017 abstract

Van Droogenbroeck, B., Kyndt, T., Romeijn-Peeters, E., Van Thuyne, W., Goetghebeur, P., Romero-Motochi, J.P., Gheysen, G. 2006. Evidence of natural hybridization and introgression between Vasconcellea species (Caricaceae) from Southern Ecuador revealed by chloroplast, mitochondrial and nuclear DNA markers. Annals of Botany 97(5):793-805. doi: 10.1093/aob/mcl038 open access

Kyndt, T., Romeijn-Peeters, E., Van Droogenbroeck, B., Romero-Motochi, J.P., Gheysen, G., Goetghebeur, P. 2005b. Species relationships in the genus Vasconcellea (Caricaceae) based on molecular and morphological evidence. Am. J. Bot. 92(6): 1033-1044. doi: 10.3732/ajb.92.6.1033 open access

Olson, M.E. 2002. Intergeneric relationships within the Caricaceae-Moringaceae clade (Brassicales) and potential morphological synapomorphies of the clade and its families. Int. J. Plant Sci. 163(1): 51-65. doi: 10.1086/324046 abstract

Van Droogenbroeck, B.V., Breyne, P., Goetghebeur, P., Romeijn-Peeters, E., Kyndt, T., Gheysen, G. 2002. AFLP analysis of genetic relationships among papaya and its wild relatives (Caricaceae) from Ecuador. Theor. Appl. Genet. 105(2-3): 289-297. doi: 10.1007/s00122-002-0983-4 abstract

Aradhya, M.K., Manshardt, R.M., Zee, F., Morden, C.W. 1999. A phylogenetic analysis of the genus Carica L. (Caricaceae) based on restriction fragment length variation in a cpDNA intergenic spacer region. Genet. Resour. Crop Evol. 46, 579-586. doi: 10.1023/A:1008786531609 abstract

Jobin-Decor, M.P., Graham, G.C., Henry, R.J., Drew, R.A., 1997. RAPD and isozyme analysis of genetic relationships between Carica papaya and wild relatives. Genet. Resour. Crop Evol. 44(5): 471-477. doi: 10.1023/A:1008644901727 abstract

Origin of Carica papaya

Solms-Laubach. 1889. Die Heimat und der Ursprung des kultivierten Melonenbaumes, Carica papaya L. Bot. Ztg. 47(44): 709-720. open access

Storey, W.B. 1976. Papaya, Carica papaya. In: Simmonds, N.W. (Ed.) Evolution of Crop Plants. Longman, London, pp. 21-24.

Wild form of Carica papaya

Chávez-Pesqueira, M., Suárez-Montes, P., Castillo, G., Núñez-Farfán, K. (2014) Habitat fragmentation threatens wild populations of Carica papaya (Caricaceae) in a lowland rainforest. Am. J. Bot., 101: 1092-1101. doi:10.3732/ajb.1400051 abstract

Brown, J.E., Bauman, J.M., Lawrie, J.F., Rocha, O.J., Moore, R.C. 2012. The structure of morphological and genetic diversity in natural populations of Carica papaya (Caricaceae) in Costa Rica. Biotropica 44, 179-188. doi: 10.1111/j.1744-7429.2011.00779.x abstract

Coppens d'Eeckenbrugge, G., Restrepo, M.T., Jiménez, D. 2007. Morphological and isozyme characterization of common papaya in Costa Rica. Acta Hort. 740, 109-120.abstract

Morshidi, M., Manshardt, R.M., Zee, F. 1995. Isozyme variability in wild and cultivated Carica papaya. HortScience 30, 809. abstract

Paz, L., Vázquez-Yanes, C. 1998. Comparative seed ecophysiology of wild and cultivated Carica papaya trees from a tropical rain forest region in Mexico. Tree Physiol. 18, 277-280. open access

Sexual systems

Aguirre, A., Vallejo-Marín, M., Piedra-Malagón, E.M., Cruz-Ortega, R., Dirzo, R. 2009. Morphological variation in the flowers of Jacaratia mexicana A. DC. (Caricaceae), a subdioecious tree. Plant Biol. (Stuttg). 11, 417-24. doi: 10.1111/j.1438-8677.2008.00154.x abstract

Aguirre, A., Vallejo-Marín, M., Salazar-Goroztieta, L., Arias, D.M., Dirzo, R. 2007. Variation in sexual expression in Jacaratia mexicana (Caricaceae) in Southern Mexico: frequency and relative seed performance of fruit-producing males. Biotropica 39, 79-86. doi: 10.1111/j.1744-7429.2006.00230.x abstract

Renner, S.S. 2004. Dioecy database open access

Renner, S.S., Ricklefs, R.E. 1995. Dioecy and its correlates in the flowering plants. Am. J. Bot. 82, 596. abstract

Storey, W.B. 1941. The botany and sex relationships of the papaya. Hawaii Agr. Exp. Sta. Bull. 87: 5-22.


Caetano, C.M., Burbano, T.C.L., Sierra, C.L.S., Tique, C.A.P., Nunes, D.G.C. 2008. Citogenética de especies de Vasconcellea (Caricaceae). Acta Agron (Palmira) 57, 241-245. open access

Costa, F.R., Pereira, T.N.S., Hodnett, G.L., Pereira, M.G., Stelly, D.M. 2008. Fluorescent in situ hybridization of 18S and 5S rDNA in papaya ( Carica papaya L.) and wild relatives. Caryologia 61, 411-416. open acces

Damasceno, C., Pedro, J., Rabelo, F., Santana, T.N., Neto, M.F., Pereira, M.G. 2009. Karyotype determination in three Caricaceae species emphasizing the cultivated form ( C . papaya L.). Caryologia 62, 10-15. open access

Kumar, L.S.S., Srinivasan, V.K. 1944. Chromosome number of Carica dodecaphylla Vell. Curr. Sci. 13: 15

Silva, E.N. da, Neto, M.F., Pereira, T.N.S., Pereira, M.G. 2012. Meiotic behavior of wild Caricaceae species potentially suitable for papaya improvement. Crop Breed. Appl. Biotechn. 12, 52-59. open access

Sex Chromosomes

Wang, J., Na, J., Yu, Q., Gschwend, A.R., Han, J., Zeng, F. 2012. Sequencing papaya X and Y h chromosomes reveals molecular basis of incipient sex chromosome evolution. doi: 10.1073/pnas.1207833109 (abstract)

Ming, R., Bendahmane, A., Renner, S.S. 2011. Sex chromosomes in land plants. Annu. Rev. Plant Biol. 62, 485-514. doi: 10.1146/annurev-arplant-042110-103914 open access

Wu, X., Wang, J., Na, J.K., Yu, Q., Moore, R.C., Zee, F., Huber, S.C., Ming, R. 2010. The origin of the non-recombining region of sex chromosomes in Carica and Vasconcellea. Plant J. 63, 801-810. doi: 10.1111/j.1365-313X.2010.04284.x open access

Ming, R., Yu, Q., Moore, P.H. 2007. Sex determination in papaya. Semin. Cell Dev. Biol. 18, 401-8. doi: 10.1016/j.semcdb.2006.11.013 (Abstract)

Liu, Z., Moore, P.H., Ma, H., Ackerman, C.M., Ragiba, M., Yu, Q., Pearl, H.M., Kim, M.S.,Charlton, J.W., Stiles, J.I., Zee, F.T., Paterson, A.H., Ming, R. 2004. A primitive Y chromosome in papaya marks incipient sex chromosome evolution. Nature 427, 348-352. abstract

Horovitz, S. 1954. Determinacion del sexo en Carica papaya L. estructura hipotetica de los cromosomas sexuales. Agronomia Tropical 3(4):229-249.

Storey, W.B. 1953. Genetics of the papaya. J. of Hered. 44, 70-78. download pdf


Ming, R., Moore, P.H. (Eds.) 2014. Genetics and Genomics of Papaya. Springer New York, New York, NY, 438 pp. Buy book

Gschwend, A.R., Wai, C.M., Zee, F., Arumuganathan, A.K., Ming, R. 2012. Genome size variation among sex types in dioecious and trioecious Caricaceae species. Euphytica 189, 461-469 abstract

Ming, R., Hou, S., Feng, Y. et al. 2008. The draft genome of the transgenic tropical fruit tree papaya (Carica papaya Linnaeus). Nature 452, 991-996. doi: doi: 10.1038/nature06856. open access


d'Eeckenbrugge, G.C., Drew, R., Kyndt, T., Scheldeman, X. 2014. Vasconcellea for Papaya Improvement, in: Ming, R., Moore, P.H. (Eds.), Genetics and Genomics of Papaya. Springer New York, New York, NY, pp. 47-79. doi: 10.1007/978-1-4614-8087-7 Buy book

Scheldeman, X., Kyndt, T., Coppens dEeckenbrugge, G.C., Ming, R., Drew, R., Van Droogenbroeck, B.V., Van Damme, P., Moore, P.H. 2011. Vasconcellea. In: Kole, C. (Ed.), Wild Crop Relatives: Genomic and Breeding Resources. Tropical and Subtropical Fruits. Springer Verlag, Berlin Heidelberg. doi: 10.1007/978-3-642-20447-0_11 Buy book

Siar, S.V., Beligan, G.A., Sajise, A.J.C., Villegas, V.N., Drew, R.A. 2011. Papaya ringspot virus resistance in Carica papaya via introgression from Vasconcellea quercifolia. Euphytica 181, 159-168. doi: 10.1007/s10681-011-0388-z abstract

Dinesh, M., Rekha, A., Ravishankar, K., Praveen, K., Santosh, L. 2007. Breaking the intergeneric crossing barrier in papaya using sucrose treatment. Sci. Hortic. 114, 33-36. doi: 10.1016/j.scienta.2007.05.010 abstract

Drew, R.A., Magdalita, P.M., O'Brien, C.M. 1998. Development of Carica interspecific hybrids. Acta Hort. 461, 285-292. abstract.

Magdalita P.M., Adkins S.W., Godwin I.D., Drew R.A. 1996. An Improved Embryo-Rescue Protocol for a Carica Interspecific Hybrid . Australian Journal of Botany 44 , 343?353. doi: 10.1071/BT9960343 abstract.

Manshardt, R.M., Zee, F.T.P., 1994. Papaya germplasm and breeding in Hawaii. Fruit Var. J. 48, 146-152. abstract

Horovitz, S., Jiménez, H. 1972. The ambisexual form of Carica pubescens Lenné et Koch analized in interspecific. Agron. Trop. 22, 475-482. open access

Horovitz, S., Jimenez, H., 1967. Cruzamientos interspecíficos e intergenéricos en Caricaceas y sus implicaciones fitotécnicas. Agron. Trop. (Maracay) 17, 323-343. open access

Sawant, A.C., 1958. Crossing relationships in the genus Carica. Evolution 12, 263-266. abstract

Agronomy, Physiology, and Chemistry

Manshardt, R.M., 1992. Papaya. In: Hammerschlag, F.A., Litz, R.E. (Eds.) Biotechnology of Perennial Fruit Crops. Cambridge University Press, Oxford, pp. 489-511

Tokuhisa, D., Cunha, D., Dos, F., Dias, S., Alvarenga, E.M., Hilst, P.C., Demuner, A.J., 2007. Compostos fenólicos inibidores da germinação de sementes de mamão (Carica papaya L.). Rev. Bras. Sementes 29, 161-168. open access

De Feo, V., De Simone, F., Arroyo, G.A., Senatore, F., 1999. Carica candicans Gray (Mito), an alimentary resource from Peruvian flora. J. Agric. Chem. 47, 3682-3684.

Willingham, B.C., White, G.A., 1976. Agronomic evaluation of prospective new crop species V. Jarilla chocola: a proteinase source. Econ. Bot. 30, 189-192. abstract

Dated phylogenies (only studies including Caricaceae)

Carvalho, F. and Renner, S.S. 2012. A dated phylogeny of the papaya family (Caricaceae) reveals the crop's closest relatives and the family's biogeographic history. Mol. Phylogenet. Evol. 65(1): 46-53. doi: 10.1016/j.ympev.2012.05.019 open access

Beilstein, M.A., Nagalingum, N.S., Clements, M.D., Manchester, S.R., Mathews, S. 2010. Dated molecular phylogenies indicate a Miocene origin for Arabidopsis thaliana. Proc. Natl. Acad. Sci. U. S. A. 107, 18724-8. open access

Wikström, N., Savolainen, V., Chase, M.W. 2001. Evolution of the angiosperms: calibrating the family tree. Proc. R. Soc. B 268, 2211-20. doi: 10.1098/rspb.2001.1782 abtract

Bell, C.D., Soltis, D.E., Soltis, P.S. 2010. The age and diversification of the angiosperms re-revisited. Am. J. Bot. 97, 1296-1303. doi: 10.3732/ajb.0900346 open access

Crop domestication

Colunga-GarcíaMarín, P., Zizumbo-Villarreal, D. 2004. Domestication of plants in Maya lowlands. Econ. Bot. 58, S101-S110.abstract

Pope, K.O., Pohl, M.E., Jones, J.G., Lentz, D.L., von Nagy, C., Vega, F.J., Quitmyer, I.R., 2001. Origin and environmental setting of ancient agriculture in the lowlands of Mesoamerica. Science 292, 1370-3. doi: 10.1126/science.292.5520.1370 abstract

Pohl, M.D., Pope, K.O., Jones, J.G., Jacob, J.S., Piperno, D.R., Susan, D., Lentz, D.L., Gifford, J.A., Danforth, M.E., Josserand, J.K., Latin, S., Antiquity, A., Dec, N., Pope, K. 1996. Early Agriculture in the Maya Lowlands. Lat. Am. Antiq. 7, 355-372. abstract

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