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Speciation in the Hawaiian "Drosophila": Sexual Selection Appears to Play an Important Role Author(S): Kenneth Y

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Speciation in the Hawaiian Speciation in the Hawaiian "Drosophila": Sexual Selection Appears to Play an Important Role Author(s): Kenneth Y. Kaneshiro Reviewed work(s): Source: BioScience, Vol. 38, No. 4, Hawaii's Unique Biology (Apr., 1988), pp. 258-263 Published by: University of California Press on behalf of the American Institute of Biological Sciences Stable URL: http://www.jstor.org/stable/1310849 . Accessed: 19/09/2012 21:35 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. University of California Press and American Institute of Biological Sciences are collaborating with JSTOR to digitize, preserve and extend access to BioScience. http://www.jstor.org Speciationin the Hawaiian Drosophila Sexual selection appears to play an important role Kenneth Y. Kaneshiro t has long been recognized that the Hawaiian drosophilids included the Hawaiian Islands possess one With their often bizarre 400 species. Since then, 89 additional of the most remarkable insect fau- species have been named and de- nas on Earth and are indeed the ideal courtship patterns, scribed (Hardy and Kaneshiro 1981). natural laboratories for the study of are At least 200-250 more new species in the processes of speciation and evolu- Hawaiian Drosophila this family are already present in the tion (Hardy and Kaneshiro 1981). the of collection at the University of Hawaii Nowhere else in the world do we find birds-of-paradise but have not been described. More so many examples of explosive adap- the insect world new species are still being collected as tive radiation-the evolution of a additional sites within the islands are large number of divergent forms from sampled, and the total Hawaiian dro- a primitive ancestor. By far the most tionary phenomena. sophilid fauna may exceed 800 outstanding of these examples is the The Hawaiian drosophilids are species. Hawaiian Drosophilidae (see reviews closely associated with vegetation in The Hawaiian drosophilids have in Carson et al. 1970, Carson and the native ecosystem. Many of the not only developed a fantastic num- Kaneshiro 1976, Hardy and Kane- species are composed of extremely ber of species, but also exhibit greater shiro 1981, Kaneshiro 1983). small populations with limited distri- diversity of forms and habits than any Here, in this isolated group of Pa- bution. While it is not known how other group in this family. The bodies cific Basin islands situated more than many species might have been lost and wings are often ornately pat- 3500 kilometers from the nearest due to perturbation of native habi- terned, and unusual modifications of land mass, are found nearly one- tats, there are clear indications of the the mouthparts and legs of the males fourth of the known species in the fragility of these endemic flies. Within frequently occur (Figure 1). Many of family Drosophilidae. More than the last two decades of intensive evo- these features would be considered to 95% (484 out of 509 species in two lutionary research on these flies, sev- signify different genera among species genera) are endemic to Hawaii; they eral species are known to have be- from other areas of the world, but in are found nowhere else in the world. come extremely rare or extinct. A the Hawaiian drosophilids, these fea- Considering Hawaii's small land number of proposed and ongoing tures have been shown to be of only mass-approximately 15,500 square projects could have a major impact species group importance. Spieth kilometers-and the geological new- on a number of these rare and threat- (1966, 1974) demonstrated that these ness of the present-day high islands of ened drosophilid species. These pro- structural peculiarities in the Hawai- the Hawaiian Archipelago, the Ha- jects include the geothermal develop- ian males are manifestations of often- waiian Drosophilidae represents one ment programs within native rain times bizarre courtship patterns. The of the most striking examples of forests on the islands of Hawaii and Hawaiian drosophilids are the birds- adaptive radiation known for any Maui and a proposal to eradicate of-paradise of the insect world. group of animals anywhere, and these tephritid fruit fly species from the With the exception of the endemic islands offer a unique opportunity for State of Hawaii by applying chemical genus Titanochaeta, which preys investigating dynamic stages of evolu- pesticides, even in areas bordering upon spider eggs, the Hawaiian dro- native habitats. It will be important to sophilids are predominantly sapro- de- The larvae feed as KennethY. Kaneshirois the Director of pay close attention to potentially phytic. scavengers and occa- the Hawaiian Pro- structive projects in order to minimize on decaying bark, leaves, Evolutionary Biology on flowers and fruits of na- at the University of Hawaii in their impact on Hawaii's extremely sionally gram tive also feed on Honolulu 96822. ? 1988 American Insti- fragile ecosystems. plants. They slime tute of Biological Sciences. Hardy's (1965) taxonomic study of fluxes and fleshy fungi. 258 BioScience Vol. 38 No. 4 Heed (1968) lists nine major ovipo- sitional substrates for these flies. Al- '/ though a few species are generalists in their ovipositional behavior (e.g., Drosophila grimshawi from the Maui complex of islands will oviposit in more than ten different families of b native plants), the majority are strict r. specialists, ovipositing in one or two plant species. It seems that, at least for the picture-winged species group of Hawaiian Drosophila, the degree of specialism in larval breeding site is associated with ovipositional behav- dri ior of the females rather than the nutritional requirements of the larval rf stages (Carson and Kaneshiro 1976). After a small piece of their natural ,ii substrate stimulates gravid females of e / specialist species to oviposit, the lar- vae will develop fully in standard laboratory medium. g In spite of their morphological, ecological, and behavioral diversities, the Hawaiian drosophilids constitute an extremely close-knit evolutionary group. From the data to date, it ap- pears that the entire endemic fauna could have arisen from a single suc- cessful founder. Although there are two major lineages in Hawaii, the drosophiloids and the scaptomyzoids, several lines of evidence show distinct similarities in the two groups, and only in Hawaii do these two groups intergrade. Throckmorton (1966, p. 386), on the basis of comparative anatomy of internal structures of the two groups, states ".. Hawaii must be considered to be the only place in the world where the otherwise sharp distinctions between Scaptomyza and tend to In a Drosophila disappear." 1. Hawaiian Drosophila species exhibit great diversity in legs, mouthparts, review of the Figure major family Drosophi- heads, and wing patterns,a. Front tarsus of D. pectinitarsus(split-tarsus subgroup). lidae, Throckmorton (1975, pp. 455- b. Fronttarsus of D. expansa (bristle-tarsussubgroup). c. Fronttibia and tarsusof D. 456) further states that although "... dasycnemia (spoon-tarsussubgroup). d. D. lasiopoda (picture-winggroup), e. D. the problem of the origin of the dro- grimshawi (picture-winggroup). f. D. aethostoma (modified-mouthpartsgroup). sophiloid and scaptomyzoid lineages g. Modifiedhead of D. heteroneura.From Hardy and Kaneshiro(1981). of Hawaii remain unresolved ... the parsimonious inference from the evi- dence, in its present state, still favors the origin of all the Hawaiian Droso- biochemical studies). a variety of genetic techniques, in- philids from a single introduction." One of the most comprehensive cluding isozyme analyses, DNA se- The ultimate goal of the systematist methods of zoological classification quencing, chromosomal analyses, hy- is to be able to arrange any group of has been developed during the last 25 bridization, and behavioral genetics, organisms in a phylogenetic scheme years by the Hawaiian Drosophila to obtain important information on that portrays evolutionary history. It Research Project, with support from the mechanism of speciation in this is clear that such phylogenetic rela- the National Institutes of Health and group of insects. tionships must be based upon sup- the National Science Foundation. In- Nevertheless, some important portive information from many disci- vestigators in Hawaii and in labora- questions remain. Did one introduc- plines (e.g., morphology, genetics, tories in the continental United States, tion into the Hawaiian Islands really droso- ecology, behavior, physiology, and Europe, Asia, and Australia are using give rise to this large, endemic 259 April 1988 philid fauna? Which particular non- The chromosome studies on the fixed inversion differences in their Hawaiian group is the ancestor of the picture-winged species of Hawaiian banding sequence, than they are to Hawaiian drosophilids? How old is Drosophila conducted by Hampton the others in the homosequential the endemic drosophilid fauna (see L. Carson and his colleagues (Carson group. On the other hand, D. vesci- Beverley and Wilson 1985)? How and Kaneshiro 1976, Carson and seta, is closely related morphological- many drosophilid species (extant and
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