AbstractThe Hawaiian Drosophilidae is one of the best examples of rapid speciation in nature. Nearly 1,000 species of endemic drosophilids have evolved in situ in Hawaii since a single colonist arrived over 25 million years ago. A number of mechanisms, including ecological adaptation, sexual selection, and geographic isolation, have been proposed to explain the evolution of this hyperdiverse group of species. Here, we examine the known ecological associations of 326 species of endemic Hawaiian Drosophilidae in light of the phylogenetic relationships of these species. Our analysis suggests that the long-accepted belief of strict ecological specialization in this group does not hold for all taxa. While many species have a primary host plant family, females will also oviposit on non-preferred host plant taxa. Host shifting is fairly common in some groups, especially the grimshawi and modified mouthparts species groups of Drosophila, and the Scaptomyza subgenus Elmomyza. Associations with types of substrates (bark, leaves, flowers) are more evolutionarily conserved than associations with host plant families. These data not only give us insight into the role ecology has played in the evolution of this large group, but can help in making decisions about the management of rare and endangered host plants and the insects that rely upon them for survival.
Ashburner, M. (1981) Entomophagous and other bizarre Drosophilidae. In: Ashburner, M., Carson, H.L. & Thompson, J.N., Jr. (Eds.) The Genetics and Biology of Drosophila, Vol. 3a. Academic Press, London, pp. 395–429.
Barker, J.S.F. Starmer, W.T. & MacIntyre, R.J. (1990) Ecological and Evolutionary Genetics of Drosophila. New York, Plenum, 542 pp.
Becerra, J.X. (2003) Synchronous coadaptation in an ancient case of herbivory. Proceedings of the National Academy of Sciences USA, 100, 12804–12807.
Beverley, S.M. & Wilson, A.C. (1985) Ancient origin for Hawaiian Drosophilidae inferred from protein comparisons. Proceedings of the National Academy of Sciences USA, 82, 4753–4757.
Bonacum, J. (2001) Molecular systematics of the Hawaiian Drosophilidae. Ph.D. Thesis Department of Biology. Yale University, New Haven.
Bonacum, J., O’Grady, P.M., Kambysellis, M.P. & DeSalle, R. (2005) Phylogeny and age of diversification of the planitibia species group of the Hawaiian Drosophila. Molecular Phylogenetics and Evolution, 37, 73–82.
Bryan, E.H. (1934) A review of the Hawaiian Diptera, with descriptions of new species. Proceedings of the Hawaiian Entomological Society, 8, 434–440 and 456–457.
Bryan, E.H. (1938) Key to the Hawaiian Drosophilidae and descriptions of new species. Proceedings of the Hawaiian Entomological Society, 10, 25–42.
Carlquist, S.H. (1980) Hawaii: A Natural History. National Tropical Botanical Garden, Lawai, Hawaii, 484 pp.
Carson, H.L. (1965) Chromosomal morphism in geographically widespread species of Drosophila. In: Baker, H.G. & Ledyard Stebbins, G. (Eds.) The Genetics of Colonizing Species. Academic Press, New York, pp. 503–531.
Carson, H.L. (1966) Chromosomal races of Drosophila crucigera from the islands of Oahu and Kauai, State of Hawaii. University of Texas Publications, 6615, 405–412.
Carson, H.L. (1986) Drosophila populations in the Ola‘a Tract, Hawaii Volcanoes National Park, 1971–1986, In: Sixth Conference in Natural Sciences, Hawaii Volcanoes National Park. Cooperative National Park Resources Studies Unit, Department of Botany, University of Hawaii at Manoa, Honolulu. pp. 3–9.
Carson, H.L. & Kaneshiro, K.Y. (1976) Drosophila of Hawaii: systematics and ecological genetics. Annual Review of Ecology and Systematics, 7, 311–345.
Clague, D.A. (1996) The growth and subsidence of the Hawaiian–Emperor volcanic chain. In: Keast, A. & Miller, S.E. (Eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam, pp. 35–50.
Daly, H.V. & Magnacca, K.N. (2003) Insects of Hawaii vol. 17. Hawaiian Hylaeus (Nesoprosopis) Bees (Hymenoptera: Apoidea). University of Hawaii Press, Honolulu, 234 pp.
DeSalle, R. (1992) The origin and possible time of divergence of the Hawaiian Drosophilidae: evidence from DNA sequences. Molecular Biology and Evolution, 9, 905–916.
Eliasson, U.H. (2004) The evolutionary patterns of the plant family Amaranthaceae on the Galpagos and Hawaiian Islands. Journal of the Torrey Botanical Society, 131, 105–109.
Fogleman, J.C., Heed, W.B. & Kircher, H.W. (1982) Drosophila mettleri and senita cactus alkaloids: fitness measurements and their ecological significance. Comparative Biochemistry and Physiology. A, Comparative Physiology, 71, 413–417.
Foote, D. & Carson, H.L. (1995) Drosophila as monitors of change in Hawaiian ecosystems. In: LaRoe, E.T., Farris, G.S., Puckett, C.E., Doran, P.D. & Mac, M.J. (Eds.) Our Living Resources: A Report to the Nation on the Distribution, Abundance, and Health of U.S. Plants, Animals, and Ecosystems. U.S. Department of the Interior, National Biological Service, Washington, D.C, pp. 368–372.
Ford, E.B. (1964) Ecological Genetics (4th edition, 1975). Chapman and Hall, London, 442 pp.
Givnish, T.J., Knox, E., Patterson, T.B., Hapeman, J.R., Palmer, J.D. & Sytsma, K.J. (1996) The Hawaiian lobelioids are monophyletic and underwent a rapid initial radiation roughly 15 million years ago [abstract]. American Journal of Botany, 83(suppl.), 159.
Hardy, D.E. (1965) Insects of Hawaii vol. 12. Diptera: Cyclorrhapha II, Series Schizophora, Section Acalypterae I. Family Drosophilidae. University of Hawaii Press, Honolulu, 814 pp.
Hardy, D.E. (1966) Descriptions and notes on Hawaiian Drosophilidae (Diptera). University of Texas Publications, 6615, 195–244.
Hardy, D.E. (1977) Review of the Hawaiian Drosophila (Antopocerus). Proceedings of the Entomological Society of Washington, 79, 82–95.
Hardy, D.E. & Kaneshiro, K.Y. (1975) Studies in Hawaiian Drosophila, miscellaneous new species, no. 1. Proceedings of the Hawaiian Entomological Society, 22, 57–64.
Hardy, D.E. & Kaneshiro, K.Y. (1979) A review of the modified tarsus species group of Hawaiian Drosophila (Drosophilidae: Diptera). Proceedings of the Hawaiian Entomological Society, 23, 71–90.
Hardy, D.E., Kaneshiro, K.Y., Val, F.C. & O’Grady, P.M. (2001) Review of the haleakalae species group of Hawaiian Drosophila (Diptera: Drosophilidae). Bishop Museum Bulletin in Entomology, 9, 1–88.
Heed, W.B. (1968) Ecology of the Hawaiian Drosophilidae. University of Texas Publications, 6816, 387–419.
Heed, W.B. (1971) Host plant specificity and speciation in Hawaiian Drosophila. Taxon, 20, 115–121.
Hollocher, H. & Templeton, A.R. (1994) The molecular through ecological genetics of abnormal abdomen in Drosophila mercatorum. VI. The non-neutrality of the Y chromosome rDNA polymorphism. Genetics, 136, 1373–1384.
ICZN. (1999) International Code of Zoological Nomenclature, Fourth Edition. 306 pp.
James, A.C., Jakubczak, J., Riley, M.P. & Jaenike, J. (1988) On the causes of monophagy in Drosophila quinaria. Evolution, 42, 626–630.
Jones, C.D. (1998) The genetic basis of Drosophila sechellia’s resistance to a host plant toxin. Genetics, 149, 1899–1908.
Jones, C.D. (2001) The genetic basis of larval resistance to a host plant toxin in Drosophila sechellia. Genetical Research, 78, 225–233.
Kam, M.W.Y. (1978) The biosystematics of the mimica subgroup of the modified mouthparts species group of Hawaiian Drosophila. Ph.D. Thesis University of Hawaii at Manoa, Honolulu.
Kambysellis, M.P. & Heed, W.B. (1971) Studies of oogenesis in natural populations of Drosophilidae. American Naturalist, 105, 31–49.
Kambysellis, M.P., Ho, K.-F., Craddock, E.M., Piano, F., Parisi, M. & Cohen, J. (1995) Pattern of ecological shifts in the diversification of Hawaiian Drosophila inferred from a molecular phylogeny. Current Biology, 5, 1129–1139.
Kaneshiro, K.Y. (1976) A revision of the generic concepts in the biosystematics of Hawaiian Drosophilidae. Proceedings of the Hawaiian Entomological Society, 22, 255–278.
Kaneshiro, K.Y. (1997) R.C.L. Perkins’ legacy to evolutionary research on Hawaiian Drosophilidae (Diptera). Pacific Science, 51, 450–461.
Kaneshiro, K.Y., Carson, H.L., Clayton, F.E. & Heed, W.B. (1973) Niche separation in a pair of homosequential Drosophila species from the island of Hawaii. American Naturalist, 107, 766–774.
Kibota, T.T. & Courtney, S.P. (1991) Jack of one trade, master of none: host choice by Drosophila magnaquinaria. Oecologia, 86, 251–260.
Kircher, H.W. (1969) Sterols in the leaves of the Cheirodendron gaudichaudii tree and their relationship to Hawaiian Drosophila ecology. Journal of Insect Physiology, 15, 1167–1173.
Kircher, H.W. & Heed, W.B. (1970) Phytochemistry and Drosophila-plant relations. Advances in Phytochemistry, 3, 191–209.
Kircher, H.W., Heed, W.B., Russell, J.S. & Grove, J. (1967) Senita cactus alkaloids: their significance to Sonoran Desert Drosophila ecology. Journal of Insect Physiology, 13, 1869–1874.
Lachance, M.-A., Starmer, W.T. & Bowles, J.M. (1989) The yeast community of morning glory and associated drosophilids in a Hawaiian kipuka. Yeast, 5, 501–504.
Lammers, T.G. (1991) Systematics of Clermontia (Campanulaceae: Lobelioideae). Systematic Botany Monographs, 32, 1–94.
Magnacca, K.N. & O’Grady, P.M. (2007) A subgroup structure for the modified mouthparts species group of Hawaiian Drosophila. Proceedings of the Hawaiian Entomological Society, 38, 87–101.
Magnacca, K.N. & O’Grady, P.M. (in press) Revision of the nudidrosophila and ateledrosophila species group of Hawaiian Drosophila (Diptera: Drosophilidae), with descriptions of twenty-two new species. Systematic Entomology.
Magnacca, K.N. & P.M. O’Grady. (submitted) Revision of the modified mouthparts species group of Hawaiian Drosophila (Diptera: Drosophilidae). III: New species in the adventitia, ceratostoma, freycinetiae, mitchelli, and setiger subgroups.
Mangan, R.L. (1978) Competitive interactions among host plant specific Drosophila species, Ph.D. Thesis. University of Arizona, Tucson.
Markow, T.A. & O’Grady, P.M. (2005) Evolutionary genetics of reproductive behavior in Drosophila: connecting the dots. Annual Review of Genetics, 39, 263–291.
Markow, T.A. & O’Grady, P.M. (2006) Drosophila: A Guide to Species Identification and Use. Academic Press, London, 500 pp.
Montgomery, S.L. (1975) Comparative breeding site ecology and the adaptive radiation of picture-winged Drosophila (Diptera: Drosophilidae) in Hawaii. Proceedings of the Hawaiian Entomological Society, 22, 65–103.
O’Grady, P.M. (2002) Species to genera: phylogenetic inference in the Hawaiian Drosophilidae. In: DeSalle, R., Giribet, G. & Wheeler, W. (Eds.) Molecular Systematics and Evolution: Theory and Practice. Birkhauser Verlag, Berlin, pp. 17–30.
O’Grady, P.M., Kam, M.W.Y., Val, F.C. & Perreira, W.D. (2003) Revision of the Drosophila mimica subgroup, with descriptions of ten new species. Annals of the Entomological Society of America, 96, 12–38.
O’Grady, P.M. and R. DeSalle. in press. Out of Hawaii: the biogeographic history of the genus Scaptomyza (Diptera: Drosophilidae). Biology Letters.
O’Grady, P.M., Val, F.C., Hardy, D.E. & Kaneshiro, K.Y. (2001) The rustica species group of Hawaiian Drosophila (Diptera: Drosophilidae). Pan-Pacific Entomologist, 77, 254–260.
Perreira, W.D. & Kaneshiro, K.Y. (1990) Three new species of picture-winged Drosophila from the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society, 30, 79–84.
Posada, D. & K.A. Crandall. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics 14, 817-818.
Pratt, L.W. & Abbott, L.L. (1997) Rare plants within managed units of Olaa Forest, Hawaii Volcanoes National Park. Cooperative National Park Resources Studies Unit Technical Report 115. Department of Botany, University of Hawaii at Manoa, Honolulu
Price, J.P. & Clague, D.A. (2002) How old is the Hawaiian biota? Geology and phylogeny suggest recent divergence. Proceedings of the Royal Society of London: Series B, Biological Sciences, 269, 2429–2435.
Robertson, F.W., Shook, M., Takei, G. & Gaines, H. (1968) Observations on the biology and nutrition of Drosophila disticha, Hardy, an indigenous Hawaiian species. University of Texas Publications, 6818, 279–299.
Ronquist, F. and Huelsenbeck, J.P.. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17, 754-755.
Russo, C.A.M., Takezaki, N. & Nei, M. (1995) Molecular phylogeny and divergence times of drosophilid species. Molecular Biology and Evolution, 12, 391–404.
Sanderson, M.J. 2003. r8s: inferring absolute rates of evolution and divergence times in the absence of a molecular clock. Bioinformatics 19, 301-302.
Scheuer, P.J. (1955) The constituents of Mokihana (Pelea anisata Mann). Chemistry and Industry, 33, 1257.
Scheuer, P.J. & Hudgins, W.R. (1964) Major constituents of the essential oil of Pelea christophersenii. Perfumery and Essential Oil Record, 55, 723–724.
Spieth, H.T. (1966) Courtship behavior of endemic Hawaiian Drosophila. University of Texas Publications, 6615, 245–313.
Spieth, H.T. (1980) Hawaiian Drosophila Project. Proceedings of the Hawaiian Entomological Society, 23, 275–291.
Starmer, W.T. (1981) A comparison of Drosophila habitats according to the physiological attributes of the associated yeast communities. Evolution, 35, 38–52.
Swezey, O.H. (1929) Notes on the egg-parasites of insects in Hawaii. Proceedings of the Hawaiian Entomological Society, 7, 282–292.
Swezey, O.H. (1934) Records of immigrant insects for 1933. Proceedings of the Hawaiian Entomological Society, 8, 544.
Swezey, O.H. (1952) Note: Insects from decaying blossoms. Proceedings of the Hawaiian Entomological Society, 14, 357.
Thomas, R.H. & Hunt, J.A. (1993) Phylogenetic relationships in Drosophila: a conflict between molecular and morphological data. Molecular Biology and Evolution, 10, 362.
Throckmorton, L.H. (1966) The relationships of the endemic Hawaiian Drosophilidae. University of Texas Publications, 6615, 335–396.
U.S. Fish and Wildlife Service. (2006) Endangered and threatened wildlife and plants; determination of status for 12 species of picture-wing flies from the Hawaiian Islands. Federal Register, 71, 26835–26852.
Wagner, W.L., Herbst, D.R. & Somer, S.H. (1999) Manual of the Flowering Plants of Hawaii, 2nd edition. University of Hawaii Press, Honolulu, 2000 pp.
Wirth, W.W. (1952) Two new spider egg predators from the Hawaiian Islands (Diptera: Drosophilidae). Proceedings of the Hawaiian Entomological Society, 14, 443–484.
Wright, S.D., Yong, C.G., Wichman, S.R., Dawson, J.W. & Gardner, R.C. (2001) Stepping stones to Hawaii: a trans-equatorial dispersal pathway for Metrosideros (Myrtaceae) inferred from nrDNA (ITS+ETS). Journal of Biogeography, 28, 769–774.