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Genetic Diversity in Eastern Polynesian Eumusa Bananas1

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Genetic Diversity in Eastern Polynesian Eumusa Bananas1 Pacific Science (1994), vol. 48, no. 1: 16-31 © 1994 by University of Hawaii Press. All rights reserved Genetic Diversity in Eastern Polynesian Eumusa Bananas1 VINCENT LEBOT, 2 BRIEN A. MEILLEUR,3 AND RICHARD M. MANSHARDT4 ABSTRACT: Genetic variation within and between the Polynesian Eumusa bananas from Hawai'i, the Marquesas, and the Society Islands is described. Morphological, isozymic, ethnographic, and linguistic-assessments ofaccessions are used to identify base clones and somatic mutants. A historical review of relevant studies is summarized. MANY KINDS OF BANANAS are cultivated on the than 70 different bananas at that time. Not­ high islands of Eastern Polynesia. Most of withstanding, after systematic collecting and these varieties are known to have been intro­ assessment, Pope (1926) recognized only 18 duced after the islands were first contacted by distinct cultivars: the undifferentiated 'Mai'a European explorers. However, well before the 'oa' clone and 17 morphotypes belonging to arrival of Europeans, bananas were brought the three principal Polynesian base clones, into Eastern Polynesia and cultivated by Poly­ iholena, maoli, andpopo'ulu (see Materials and nesians. In this paper, we describe the extent Methods section for explanation oftaxonomic of genetic variation within and between the terminology). Four additional morphotypes Polynesian Eumusa bananas from Hawai'i, of the base clones and a fifth undifferentiated the Marquesas, and the Society Islands. cultivar, 'Hiipai', were later recognized (Pope 1927) from fragmentary accounts of earlier writers, such as Thrum (1890), Higgins (1904), and MacCaughey (1918). HISTORICAL REVIEW Elsewhere in Polynesia, Brown (1931) Interest in cataloging the Polynesian claimed that "not less than 75 varieties seem bananas began in the late nineteenth century to have been cultivated by the ancient Mar­ (Thrum 1890), but the most useful work dates quesans," though he cited only 50 names, to the first halfof the twentieth century.·Pope some ofwhich clearly refer to recent introduc­ (1926), referring to a manuscript written in tions and others to Musa fehi Bert. ex Vieill. 1870, reported that the Hawaiians of Kona bananas ofthe Australimusa series. Dordillon (Hawai'i Island) knew and named no fewer (1931-1932) cited 40 banana names; as in Brown, several references were recent arrivals (e.g., kina undoubtedly refers to Chinese or 1 Funding for this research was provided by the Cavendish clones) and at least one, huetu, Hawaii-Bishop Research Institute through grants to refers to M. fehi. B.A.M. and R.M.M., and the Bishop Museum's Hermes The geographic origin ofthese bananas has Small Grants Program to B.A.M. This is Journal Series been a longstanding subject of speculation No. 3827 of the Hawaii Institute of Tropical Agriculture and Human Resources. Manuscript accepted 20 May and debate among banana taxonomists. Mac­ 1993. Daniels (1947) suggested that the Hawaiian 2CTFT/CIRAD, P.O. Box 745, Antananarivo, bananas were brought by the Polynesians Madagascar. from the Marquesas or the Society Islands. 3Center for Plant Conservation, Missouri Botanical Garden, P.O. Box 299, St. Louis MO 63166. Simmonds (1954) initially concluded that the 4 Department ofHorticulture, University ofHawai'i at Hawaiian cultivars originated in Malaya or Manoa, Honolulu, Hawai'i 96822. Indo-Malaya, from whence they were spread 16 Genetic Variations in Eumusa Bananas-LEBoT ET AL. 17 to Polynesia and Hawai'i in cultivation; later, tensively and evaluated as crop genetic re­ Simmonds (Stover & Simmonds 1987) re­ sources (Persley and De Langhe 1987). It is versed himself, stating that "these bananas are our purpose here, in part, to respond to this clearly not of Indo-Malaysian origin ... nor suggestion. can they have come from Papua New Guinea." Despite this last assertion, recent work demonstrates that the Polynesian SCOPE OF THE PROBLEM Eumusa bananas are not limited in their dis­ tribution to Eastern Polynesia and that Tremendous morphological variation ex­ Papua New Guinea is the most likely region ists within the cultivated bananas. World­ of their domestication (Horry 1989, Daniells wide, there may be as many as 500 cultivars 1990, Tezenas du Montcel 1990, Lebot et al. (Simmonds 1976, Purseglove 1988), most of 1993). Sterility in these bananas ensures that which are sterile and parthenocarpic, and they could only have come into and through therefore vegetatively propagated. Polynesia via human agency. Thus, their Most edible bananas have been domesti­ characterization as "native" in Hawai'i by cated from Musa acuminata Colla and from MacCaughey (1918) and others is inappro­ hybrids of this species with M. balbisiana priate. Colla. Traditionally, classification ofdomesti­ Several of the Polynesian cultivars have cated bananas and plantains has been accom­ been incorporated into international germ­ plished by scoring clones for 15 morphologi­ plasm collections and their agronomic per­ cal characters that differ between the parent formance evaluated. We have observed that species (Simmonds & Shepherd 1955). This they are highly susceptible to Panama dis­ method, coupled with chromosome counts, ease, a wilt caused by Fusarium oxysporum has permitted descriptions of clones in terms (Schlecht.) f.sp. cubense (E. F. Smith) Snyd. & ofploidy levels and the genomic contributions Hans.; to nematodes; and to corm borers of the two parental species. Recent analyses (Cosmopolites sp.). Polynesian bananas have of isozyme variation in banana clones have also suffered in competition with aggressive generally confirmed the relationships pro­ exotic plants and from development of a posed on the basis of morphological data commercial banana industry based on non­ (Jarret and Litz 1986a, b, Horry 1989, Lebot Polynesian cultivars. Although Polynesian et al. 1993). Those investigations demon­ bananas show some resistance to black leaf strated the existence of species-specific streak, a widespread banana disease caused by allozymes in M. acuminata and M. balbisiana, Mycosphaerella jijiensis Morelet, the com­ and multivariate analysis ofisozyme data has bined effects ofthese pests, other diseases, and produced clusters of clones that correspond economic changes are undoubtedly responsi­ neatly with genomic groupings (AA, AAA, ble for their drastic reduction and extirpation AAB, ABB, etc.) determined by morphologi­ on many Eastern Polynesian islands. Never­ cal analysis. theless, they are still occasionally found for Both of the above approaches have identi­ sale in local markets. With the exception of fied the Hawaiian base clones, maoli,popo'ulu, 'Hiipai', which is a sweet-fleshed dessert and iholena, as belonging to the AAB genomic banana, and 'Mai'a 'oa', which is seedy and group (although Stover and Simmonds [1987], inedible, all of the Polynesian Eumusa culti­ using only morphological data, misclassified vars are "cooking bananas," with the starchy iholena as a member of the AAA genomic flesh typical of plantains. Several continue to group). These three base clones of the Pacific be used sporadically in Eastern Polynesian Plantain subgroup have also been shown to medicine (Gutmanis 1976). have different and diagnostic zymotypes It has been suggested that cultivars of the (Lebot et al. 1993). However, neither of the maoli and popo'ulu base clones-still poorly above methods of classification can resolve known internationally-becollected more ex- the subtle distinctions between morphotypes 18 PACIFIC SCIENCE, Volume 48, January 1994 of the same base clone; that is differences MATERIALS AND METHODS arising by mutation that have be~n noted and named by both ancient and modern farmers. Taxonomical Terminology In Polynesia, as elsewhere in the Pacific, banana names consist either of a generic We follow the taxonomical terminology of head term meaning "banana" (e.g., mai'a in Stover and Simmonds (1987) and use the term Hawai'i and Tahiti [syn. mei'a in Tahiti] and "series" to refer to the major edible banana meika or mei'a in the Marquesas) followed by categories having basic chromosome numbers a secondary epithet that generally designates of 10 (Australimusa series) and II (Eumusa the clone (as in the Hawaiian 'Mai'a hiipai', series). "Group" is used to refer to major or 'Mai'a maoli'), or the epithet stands alone subdivisions within series. These are desig­ (as in 'Hiipai', or becomes the head term nated by letters that indicate both ploidy for.a modifier denoting minor morphological level and the genomic composition (e.g., AAB vanants or morphotypes (such as 'Maoli hai' group of the Eumusa series) with regard to or 'Iholena lele', etc.). Common qualifiers for their parent species M. acuminata (A genome) morphotypes reflect variation noted in aerial and/or M. balbisiana (B genome). "Sub­ parts of the plant, which are often related to group" refers to distinctive sets of clones distinctive pigmentation or size. For example, within groups, such as the Pacific Plantain 'ele'ele means "black," and designates a mor­ subgroup of the AAB group. Because of the photype of the Hawaiian maoli cultivar ex­ high degree of selection in some Polynesian hibiting intense black pigmentation on the Eumusa bananas, we modify Stover and Sim­ pseudoste1? and. petiole bases. Farmers may monds somewhat and use the term "clone" to refer to this cultIvar as 'Mai'a maoli 'ele'ele' distinguish bananas within subgroups that although 'Maoli 'ele'ele' or sometimes just differ in both morphological and isozymic "Ele'ele' are more common. characters, and "base
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