Abstract
The zeylanica group is one of the six species groups of the anthophilic genus Colocasiomyia de Meijere in the family Drosophilidae. In addition to two known species, five morphospecies have been recognized as members of this species group but left undescribed formally. In this study, species delimitation of these putatively new species was determined by barcoding of the mitochondrial COI (cytochrome c oxydase subunit I) gene and morphological comparison. Phylogenetic relationships within the genus Colocasiomyia were inferred by a cladistic analysis of 89 morphological characters. Based on the results of these analyses, we redefined the zeylanica species group and established two subgroups within it: the zeylanica subgroup comprised of C. zeylanica, C. nepalensis, C. pinangae sp. nov., C. besaris sp. nov. and C. luciphila sp. nov., and the oligochaeta subgroup of C. oligochaeta sp. nov. and C. grimaldii sp. nov. In addition, we briefly address the anthophilic habits of drosophilid flies using palm (Arecaceae) inflorescences, especially of the zeylanica group, compiling scattered collection records from the Oriental and Papuan regions.
References
- Atmowidi, T., Kahono, S., Dorly, Rahmawati, S.D., Reinaldo, D. & Ummah, E.S. (2021) The diversity of insect visitors on Indonesian Salacca (Salacca spp.) in Mekarsari Fruits Garden, Bogor, Indonesia. Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis, 69, 267–273. https://doi.org/10.11118/actaun.2021.024 DOI: https://doi.org/10.11118/actaun.2021.024
- Bøgh, A. (1996) The reproductive phenology and pollination biology of four Calamus (Arecaceae) species in Thailand. Principes, 40, 5−15.
- Cumming, J.M. & Wood, D.M. (2017) Adult morphology and terminology. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.), Manual of Afrotropical Diptera. Volume 1. Introductory Chapters and Keys to Diptera Families. South African National Biodiversity Institute, Pretoria, pp. 89–133.
- de Meijere, J.C.H. (1914) Studien über südostasiatische Dipteren IX. Tijdschrift voor Entomologie, 57, 137–275.
- Duda, O. (1923) Die orientalischen und australischen Drosophiliden-Arten (Dipteren) des Ungarischen National-Museums zu Budapest. Annales Historico-Naturales Musei Nationalis Hungarici, 20, 24‒59.
- Duda, O. (1924) Beitrag zur Systematik der Drosophiliden unter besonderer Berücksichtigung der paläarktischen u. orientalischen Arten (Dipteren). Wiegmann’s Archiv für Naturgeschichte, 90 (A) 3, 172–234, 7 pls.
- Essig, F.B. (1973) Pollination in some New Guinea palms. Principes, 17, 75–83.
- Fartyal, R.S., Gao, J.J., Toda, M.J., Hu, Y.G., Takano, K.T., Suwito, A., Katoh, T., Takigahira, T. & Yin, J.T. (2013) Colocasiomyia (Diptera: Drosophilidae) revised phylogenetically, with a new species group having peculiar lifecycles on monsteroid (Araceae) host plants. Systematic Entomology, 38, 763–782. https://doi.org/10.1111/syen.12027 DOI: https://doi.org/10.1111/syen.12027
- Ferguson, I.K., Dransfield, J., Page, F.C. & Thanikaimoni, G. (1983) Notes on the pollen morphology of Pinanga with special reference to P. aristata and P. pilosa (Palmae: Arecoideae). Grana, 22, 65–72. https://doi.org/10.1080/00173138309431967 DOI: https://doi.org/10.1080/00173138309431967
- Finet, C., Kassner, V.A., Carvalho, A.B., Chung, H., Day, J.P., Day, S., Delaney, E.K., De Ré, F.C., Dufour, H.D., Dupim, E., Izumitani, H.F., Gautério, T.B., Justen, J., Katoh, T., Kopp, A., Koshikawa, S., Longdon, B., Loreto, E.L., Nunes, M.D.S., Raja, K.K.B., Rebeiz, M., Ritchie, M.G., Saakyan, G., Sneddon, T., Teramoto, M., Tyukmaeva, V., Vanderlinde, T., Wey, E.E., Werner, T., Williams, T.M., Robe, L.J., Toda, M.J. & Marlétaz, F. (2021) DrosoPhyla: Resources for drosophilid phylogeny and systematics. Genome Biology and Evolution, 13, evab179. https://doi.org/10.1093/gbe/evab179 DOI: https://doi.org/10.1101/2021.03.23.436709
- Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I for diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
- Grimaldi, D.A. (1991) Systematics of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae): cladistics, a new generic synonym, new record, and a new species from Nepal. Entomologica Scandinavica, 22, 417–426. https://doi.org/10.1163/187631291X00219 DOI: https://doi.org/10.1163/187631291X00219
- Grimaldi, D., Ervik, F. & Bernal, R. (2003) Two new Neotropical genera of Drosophilidae (Diptera) visiting palm flowers. Journal of the Kansas Entomological Society, 76, 109–124. [http://www.jstor.org/stable/25086096]
- Hadley, A. (2010) Combine ZP Software. Available from: https://combinezp.software. informer.com/ (accessed 7 June 2022)
- Henderson, A. (1986) A review of pollination studies in the Palmae. Botanical Review, 52, 221–259. https://doi.org/10.1007/BF02860996 DOI: https://doi.org/10.1007/BF02860996
- Kim, B.Y., Wang, J.R., Miller, D.E., Barmina, O., Delaney, E., Thompson, A., Comeault, A.A., Peede, D., D’Agostino, E.R.R., Pelaez, J., Aguilar, J.M., Haji, D., Matsunaga, T., Armstrong, E.E., Zych, M., Ogawa, Y., Stamenković-Radak, M., Jelić, M., Veselinović, M.S., Tanasković, M., Erić, P., Gao, J.-J., Katoh, T.K., Toda, M.J., Watabe, H., Watada, M., Davis, J.S., Moyle, L.C., Manoli, G., Bertolini, E., Koštál, V., Hawley, R.S., Takahashi, A., Jones, C.D., Price, D.K., Whiteman, N., Kopp, A., Matute, D.R. & Petrov, D.A. (2021) Highly contiguous assemblies of 101 drosophilid genomes. eLife, 10, e66405. https://doi.org/10.7554/eLife.66405 DOI: https://doi.org/10.7554/eLife.66405
- Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874. https://doi.org/10.1093/molbev/msw054 DOI: https://doi.org/10.1093/molbev/msw054
- Lachaise, D. & Tsacas, L. (1983) Breeding-sites in tropical African drosophilids. In: Ashburner, M., Carson, H.L. & Thompson Jr., J.N. (Eds.), The Genetics and Biology of Drosophila. Vol. 3d. Academic Press, London, pp. 221–332.
- Li, N.N., Toda, M.J., Fu, Z., Li, S.H. & Gao, J.J. (2014) Taxonomy of the Colocasiomyia gigantea species group (Diptera, Drosophilidae), with descriptions of four new species from Yunnan, China. ZooKeys, 406, 41–64. https://doi.org/10.3897/zookeys.406.7176 DOI: https://doi.org/10.3897/zookeys.406.7176
- McQueen, E.W., Afkhami, M., Atallah, J., Belote, J.M., Gompel, N., Heifetz, Y., Kamimura, Y., Kornhauser, S.C., Masly, J.P., O’Grady, P., Peláez, J., Rebeiz, M., Rice, G., Sánchez-Herrero, E., Nunes, M.D.S., Rampasso, A.S., Schnakenberg, S.L., Siegal, M.L., Takahashi, A., Tanaka, K.M., Turetzek, N., Zelinger, E., Courtier-Orgogozo, V., Toda, M.J., Wolfner, M.F. & Yassin, A. (2022) A standardized nomenclature and atlas of the female terminalia of Drosophila melanogaster. Fly, 16, 128–151. https://doi.org/10.1080/19336934.2022.2058309 DOI: https://doi.org/10.1080/19336934.2022.2058309
- Okada, T. (1986) Taximetrical analyses of costal chaetotaxy of the genus Drosophilella Duda, with description of a new species from Sri Lanka (Diptera, Drosophilidae). Proceedings of the Japanese Society of Systematic Zoology, 34, 53‒59.
- Okada, T. (1987) Further notes on the genus Drosophilella Duda with descriptions of two new species from Indonesia (Diptera, Drosophilidae). Proceedings of the Japanese Society of Systematic Zoology, 36, 38–45.
- Okada, T. (1988) Taxonomic note on Colocasiomyia cristata de Meijere (Diptera, Drosophilidae) with generic synonymy. Proceedings of the Japanese Society of Systematic Zoology, 37, 34–39.
- Rice, G., David, J.R., Kamimura, Y., Masly, J.P., Mcgregor, A.P., Nagy, O., Noselli, S., Nunes, M.D.S., O’Grady, P., Sánchez-Herrero, E., Siegal, M.L., Toda, M.J., Rebeiz, M., Courtier-Orgogozo, V. & Yassin, A. (2019) A standardized nomenclature and atlas of the male terminalia of Drosophila melanogaster. Fly, 13, 51–64. https://doi.org/10.1080/19336934.2019.1653733 DOI: https://doi.org/10.1080/19336934.2019.1653733
- Sahbou, A.-E., Iraqi, D., Mentag, R. & Khayi, S. (2022) BuscoPhylo: A webserver for Busco-based phylogenomic analysis for non-specialists. Research Square, (preprint). https://doi.org/10.21203/rs.3.rs-1997596/v1 DOI: https://doi.org/10.21203/rs.3.rs-1997596/v1
- Shi, T., Toda, M.J., Takano, K.T., Yafuso, M., Suwito, A., Wong, S.Y., Shang, S.Q. & Gao, J.J. (2019) A review of taxonomy and flower-breeding ecology of the Colocasiomyia toshiokai species group (Diptera: Drosophilidae), with description of a new species from Indonesia. European Journal of Entomology, 116, 341–361. https://doi.org/10.14411/eje.2019.037 DOI: https://doi.org/10.14411/eje.2019.037
- Silberbauer-Gottsberger, I. (1990) Pollination and evolution in palms. Phyton (Horn, Austria), 30, 213–233.
- Sultana, F., Hu, Y.G., Toda, M.J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684–702. https://doi.org/10.1111/j.1365-3113.2006.00344.x DOI: https://doi.org/10.1111/j.1365-3113.2006.00344.x
- Sultana, F., Toda, M.J., Yafuso, M., Lakim, M.B., Mohamed, M.B. & Cuong, N.M. (2002) A new species-group of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae), with descriptions of two new species from eastern Malaysia and Vietnam. Entomological Science, 5, 305–315.
- Swofford, D.L. (2003) PAUP*: Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. Available from: https://paup.phylosolutions.com/ (accessed 11 August 2020)
- Takano, K.T., Gao, J.-J., Hu, Y.-G., Li, N.-N., Yafuso, M., Suwito, A., Repin, R., Pungga, R.A.S., Meleng, P.A., Kaliang, C.H., Chong, L. & Toda, M.J. (2021) Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species. Zootaxa, 5079 (1), 1–70. https://doi.org/10.11646/zootaxa.5079.1.1 DOI: https://doi.org/10.11646/zootaxa.5079.1.1
- Takano, K.T.., Repin, R., Mohamed, D.M.B. & Toda, M.J. (2012) Pollination mutualism between two taxonomically undescribed Colocasiomyia species (Diptera: Drosophilidae) and their host, Alocasia macrorrhizos (Araceae), in Sabah, Borneo. Plant Biology, 14, 555–564. https://doi.org/10.1111/j.1438-8677.2011.00541.x DOI: https://doi.org/10.1111/j.1438-8677.2011.00541.x
- Takano, K.T.., Suwito, A., Gao, J.J. & Yin, J.T. (2011) Molecular phylogeny of the cristata species group of the genus Colocasiomyia (Diptera: Drosophilidae). Low Temperature Science, 69, 19–28. Available from: https://eprints.lib.hokudai.ac.jp/dspace/handle/2115/45185 (Accessed 3 May 2023)
- Takenaka, K., Yin, J.T., Wen, S.Y. & Toda, M.J. (2006) Pollination mutualism between a new species of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae) and Steudnera colocasiifolia (Araceae) in Yunnan, China. Entomological Science, 9, 79–91. https://doi.org/10.1111/j.1479-8298.2006.00156.x DOI: https://doi.org/10.1111/j.1479-8298.2006.00156.x
- Toda, M.J., Takano, K.T., Katoh, T., Xiao, L., Gao, J.-J. & Yafuso, M. (2022) Coexistence mechanisms of Colocasiomyia species (Diptera: Drosophilidae) sharing inflorescences of Alocasia odora (Araceae) as a host plant: Comparison between two- and three-species systems. Entomological Science, 25, e12506. https://doi.org/10.1111/ens.12506 DOI: https://doi.org/10.1111/ens.12506
- Valadão, H., Proença, C.E., Kuhlmann, M.P., Harris, S.A. & Tidon, R. (2019) Fruit-breeding drosophilids (Diptera) in the Neotropics: playing the field and specialising in generalism? Ecological Entomology, 44, 721‒737. https://doi.org/10.1111/een.12769 DOI: https://doi.org/10.1111/een.12769
- Watrous, L.E. & Wheeler, Q.D. (1981) The outgroup comparison method of character analysis. Systematic Zoology, 30, 1−11. https://doi.org/10.1093/sysbio/30.1.1 DOI: https://doi.org/10.1093/sysbio/30.1.1
- Yafuso, M., Sultana, F., Sasaki, Y. & Toda, M.J. (2000) A new species of Colocasiomyia de Meijere (Diptera, Drosophilidae) from North Sulawesi, Indonesia. Entomological Science, 3, 115‒119.
- Yafuso, M., Toda, M.J. & Sembel, D.T. (2008) Arengomyia, new genus for the Colocasiomyia arenga species group (Diptera: Drosophilidae), with description of a new species. Entomological Science, 11, 391–400. https://doi.org/10.1111/j.1479-8298.2008.00285.x DOI: https://doi.org/10.1111/j.1479-8298.2008.00285.x
- Yassin, A. (2013) Phylogenetic classification of the Drosophilidae Rondani (Diptera): the role of morphology in the postgenomic era. Systematic Entomology, 38, 349‒364. https://doi.org/10.1111/j.1365-3113.2012.00665.x DOI: https://doi.org/10.1111/j.1365-3113.2012.00665.x
- Yassin, A. & David, J.R. (2010) Chapter 3, Phylogenetic biogeography of Afrotropical Drosophilidae. In: Gailis, M. & Kalninš, S. (Eds.), Biogeography. Nova Science Publishers, Inc., New York, New York, pp. 105‒137.
- Zhang, W.X. & Toda, M.J. (1992) A new species-subgroup of the Drosophila immigrans species-group (Diptera, Drosophilidae), with description of two new species from China and revision of taxonomic terminology. Japanese Journal of Entomology, 60, 839‒850.