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RCL Perkins' Legacy to Evolutionary Research On Pacific Science (1997), vol. 51, no. 4: 45~61 © 1997 by University of Hawai'i Press. All rights reserved R. C. L. Perkins' Legacy to Evolutionary Research on Hawaiian Drosophilidae (Diptera)l KENNETH Y. KANESHIR02 ABSTRACT: R. C. L. Perkins' influence on evolutionary research on the Hawaiian Drosophilidae is presented. His observations of the bizarre secondary sexual struc­ tures in this group led evolutionary biologists to focus research on the role of sexual selection in speciation and the evolutionary processes responsible for the proliferation of Drosophila species in the native Hawaiian fauna. A review of early taxonomic treatment ofthe group and some ofthe ecological novelties of the group are discussed. A better understanding of the genetics, ecology, behavior, morphology, etc. resulted in a revision of the generic concepts of the group, and subsequent phylogenetic studies using modem tools of molecular biology have confirmed the monophyletic relationships among the species in this group. FROM 1893 TO 1897, R. C. L. Perkins conducted the endemic species largely consisting of small an extensive survey of Hawai'i's insect fauna. Dolichopodidae, which shrink and distort on The results of Perkins' collections and the drying, and of infinite numbers of Drosophili­ detailed field notes he recorded for the speci­ dae, many ofthese also becoming distorted, have mens including host records provided the basis been little collected" (Perkins 1913). He had for his publication in the Fauna Hawaiiensis difficulty in preserving these soft-bodied insects (1913). Perkins' efforts and careful record keep­ under the harsh conditions he faced in the field ing provide extremely valuable baseline infor­ and therefore he did not make a special effort mation that we can use to infer relative to collect dipteran species as he did some of abundance ofthe populations as well as the qual­ the other major orders of insects. Nevertheless, ity of the habitat in which these species lived a based on what he saw of the drosophilid fauna century ago. It will enable us to compare esti­ in Hawai'i, he made the following comments: mates of diversity in the insect fauna with sur­ "Drosophila is represented by an assemblage of veys conducted during later decades especially species, exhibiting great diversity in structure in two groups that Perkins surveyed extensively: and appearance. ... At present these insects, the Megalagrion damselflies and the platynine many of which are obscure and minute forms, carabid beetles. have been very imperfectly collected. To make Although Perkins focused much of his efforts an approximately complete collection and thor­ in collecting many of the groups of insects in ough study of the Hawaiian species would Hawai'i, there were some groups that presented require the devotion of many years of special logistical problems for him primarily because of work. Not less than 250 species must exist in the difficult conditions fieldwork presented to the islands, and double that number may very entomologists of his time. It was clear that Per­ probably occur" (Perkins 1913:189). kins was "uncomfortable" with collecting dip­ Perkins recognized the tremendous diversity teran species in the Hawaiian fauna. In his in the Drosophilidae, and his assessment of the introduction to Fauna Hawaiiensis, he com­ group was echoed by Elwood C. Zimmerman, mented that"... the minute and obscure Diptera, another entomologist with considerable experi­ ence in the Hawaiian fauna, who wrote a pas­ 1 This is contribution no. 1996-020 of the Hawai'i Bio­ sionate plea to geneticists and evolutionists logical Survey. Manuscript accepted 3 February 1997. about the remarkable drosophilid fauna in 2 Center for Conservation Research and Training, Uni­ versity of Hawai 'i at Manoa, 3050 Maile Way, Gilmore 409, Hawai'i and the opportunity to conduct Honolulu, Hawai'i 96822. "advanced research" on this group of flies. He 450 Perkins' Legacy to Research on Drosophilid Evolution-KANEsHIRo 451 said: "Since I became aware, many years ago, sophila to justify continued funding for another of the astonishing development of Drosophila 5 yr. in Hawaii, I have tried to interest geneticists and By the summer of 1963, Stone had assembled evolutionists in the fauna, but I fear that it has a team of researchers with expertise in nearly been considered, at least by some workers, that every aspect of evolutionary biology including my descriptions of the size and diversity of the taxonomy, morphology, ecology, behavior, cyto­ fauna are exaggerated. I do not exaggerate. It is genetics, and developmental and molecular biol­ possible that the Hawaiian drosophilid fauna ogy. Seven senior scientists from across the may be the most remarkable in the world. In United States, plus Wilson Stone and Elmo Hawaii is found a range from unusually small Hardy as principal investigators, arrived in species to absolute giants up to one centimeter Hawai'i to launch a multidisciplinary research across, and there is much morphological diver­ effort in answer to Zimmerman's plea to investi­ sity. There may be as many as 300 species con­ gate the genetics and evolution of this remark­ centrated in an area smaller than the little state able fauna. That was the beginning of the so­ of Massachusetts or less than one-fifth the size called Hawaiian Drosophila Project (Spieth of Ireland. Where else has such a drosophilid 1980). Since then, with continued funding sup­ fauna developed? Is this fauna not worthy of port from the National Institutes of Health, the detailed attention by those equipped to do National Science Foundation, and other private advanced research on the genetics and evolution organizations and foundations, nearly 80 senior of this group of fascinating flies?" (Zimmer­ scientists from all over the world have traveled man 1958:557). to Hawai'i to investigate some aspect of the Thus, Perkins' early observations of the evolutionary biology of this group. In addition, Hawaiian Drosophilidae had an influence on the more than 400 undergraduate and graduate stu­ future evolutionary research of this group. In dents and postdoctoral fellows have participated 1948, D. Elmo Hardy joined the Department of in the project either as laboratory technicians or Entomology at the University of Hawai'i as a in conducting research on these flies as part of specialist on the systematics of fruit flies in the their educational curricula. family Tephritidae. Although his primary research interests were in the tephritids of the BriefHistory of the Taxonomy of the Oriental and Pacific regions, as he began to col­ Hawaiian Drosophilidae lect flies from throughout the Hawaiian Islands, he carne to the realization that the drosophilid In 1952, Wheeler listed 73 species of dro­ fauna of Hawai'i was indeed a most remarkable sophilids from the Hawaiian Islands, two of group and that there was a tremendous opportu­ which were described by R. C. L. Perkins (1913), nity to conduct intensive research on their biol­ 45 by P. H. Grimshaw (1901-1902), and the rest ogy. After Zimmerman's (1958) plea to by Knab (1914), Sturtevant (1921), Bryan (1934, geneticists and evolutionists to investigate the 1938), Malloch (1938), Zimmerman (1938), and Hawaiian fauna, Hardy visited with Wilson S. Wirth (1952). By the time of the start of the Stone, director of the Genetics Foundation at Hawaiian Drosophila Project in 1963, Hardy had the University of Texas at Austin, in 1962 and already committed nearly 10 yr to naming and discussed the possibility ofa joint research proj­ describing more than 300 new species ofHawai­ ect between the University of Hawai'i and the ian Drosophilidae and redescribing some of the University of Texas on the genetics and evolu­ species treated by his predecessors. His mono­ tion of the Hawaiian Drosophilidae. Under the graph on the systematics of this group was pub­ leadership of Stone and Hardy, proposals were lished in 1965 and listed 400 species in nine prepared and submitted to the National Institutes genera. Since then, Hardy (1966), Hardy and of Health (NIH) and the National Science Foun­ Kaneshiro (1968, 1969, 1971, 1972, 1975a,b, dation to study this group. The proposal to the 1979), Kaneshiro (l969b), and Perreira and NIH was successful in securing funds for 1 yr Kaneshiro (1990) have named and described to support a team ofresearchers to obtain enough another 111 species in the Hawaiian fauna. So, preliminary information on the Hawaiian Dro- with a current total of 511 described species in 452 PACIFIC SCIENCE, Volume 51, October 1997 the family Drosophilidae, it appears that Per­ and, in fact, could give a misleading impression kins' prediction of the number of species in this of evolutionary divergence in the Hawaiian fauna may have been almost prophetic. How­ fauna. It was shown that the so-called "key" ever, a cursory inspection of the Hawaiian dro­ generic characters were secondary sexual struc­ sophilid collection at the University of Hawai'i tures of the males that were used in the complex indicates that there are still many more unde­ courtship displays observed in these species. scribed species in the fauna. It is estimated that Females lacked these structures and for the most there may be as many as 250-300 more unde­ part could not be differentiated from typical Dro­ scribed species already in the collections and sophila species. Thus, species previously placed more new species are still being discovered as in separate genera proved to be nothing more new collecting techniques are implemented and than species groups within the genus Drosoph­ previously unsampled localities are studied. ila. The endemic genus "Idiomyia" (Grimshaw) Kaneshiro (1993) estimated that 1000 species of had already been sunk as a synonym ofDrosoph­ drosophilids may be present in the Hawaiian ila (Hardy and Kaneshiro 1968), and Kaneshiro fauna. (1976a) sank the three remaining endemic gen­ Although Hardy's (1965) treatment of the era, Antopocerus, Nudidrosophila, andAteledro­ endemic fauna included nine genera, several sophila (listed in Hardy's [1965] monograph) as lines of evidence indicated that the entire fauna synonyms of the genus Drosophila.
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