Allozyme Diversity and Lntrogression in the Galapagos Islands Endemic Gossypium Datwinii and Its Relationship to Continental G
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Botany Publication and Papers Botany 1990 Allozyme Diversity and lntrogression in the Galapagos Islands Endemic Gossypium datwinii and its Relationship to Continental G. barbadense Jonathan F. Wendel Iowa State University, [email protected] Richard G. Percy United States Department of Agriculture Follow this and additional works at: http://lib.dr.iastate.edu/bot_pubs Part of the Botany Commons, Evolution Commons, and the Molecular Genetics Commons Recommended Citation Wendel, Jonathan F. and Percy, Richard G., "Allozyme Diversity and lntrogression in the Galapagos Islands Endemic Gossypium datwinii and its Relationship to Continental G. barbadense" (1990). Botany Publication and Papers. 10. http://lib.dr.iastate.edu/bot_pubs/10 This Article is brought to you for free and open access by the Botany at Iowa State University Digital Repository. It has been accepted for inclusion in Botany Publication and Papers by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Allozyme Diversity and lntrogression in the Galapagos Islands Endemic Gossypium datwinii and its Relationship to Continental G. barbadense Abstract Gossypium darwinii Watt is a tetraploid cotton endemic to the Galapagos Islands. Opinion has been divided as to whether or not it deserves recognition at the specific ar nk, with some considering it a variety of its presumed progenitor, the widely distributed South American species G. barbadense L. A previous hypothesis states that much of the perceived intergradation between the two taxa arose as a consequence of introgression from G. barbadense following its introduction to the archipelago during the past several hundred years. We performed allozyme analysis on 58 accessions of G. darwinii from six islands, using 17 enzymes collectively encoded by 59 loci. Levels of variation were high for an island endemic, with a mean number of alleles per locus of 1.34 and an average panmictic heterozygosity of 0.062. Principal component analysis revealed clustering of accessions according to their island of origin, and a spatial pattern of island-clusters that approximates geographical relationships among islands. Genetic relationships of G. darwinii with G. barbadense and G. hirsutum L. were studied using previously generated allozyme data. Significant introgression of G. hirsutum alleles was detected; however morphological considerations support the hypothesis that much of G. darwinii's diversity stems from interspecific eg ne flow from G. barbadense, Evidence is presented suggesting that the occurrence of G. hirsutum alleles in G. darwinii derives not from direct hybridization, but from a mediated transfer through introduced, G. hirsutum-introgressed;G. barbadense. Gossypium darwinii and G. barbadense are nearly fixed for different alleles at four loci and each contains a large number of unique alleles. Notwithstanding the high interspecific eiN 's genetic identity (0.949), the allozyme data support geographical and morphological evidence in suggesting that a specific ar nk for G. darwinii is warranted. Keywords Gossypium darwinii, G. barbadense, Malvaceae, cotton, introgression, Galapagos Islands, genetic variation, island evolution, allozymes, molecular divergence Disciplines Botany | Evolution | Molecular Genetics Comments This article is from Biochemical Systematics and Ecology 18 (1990): 517, doi:10.1016/ 0305-1978(90)90123-W. Rights Works produced by employees of the U.S. Government as part of their official duties are not copyrighted within the U.S. The onc tent of this document is not copyrighted. This article is available at Iowa State University Digital Repository: http://lib.dr.iastate.edu/bot_pubs/10 Biochemical Systematics and Ecology, Vol. 18, No. 7/8, pp. 517-528, 1990. 0305-1978/90 $3.00 + 0.00 Printed in Great Britain. © 1991 Pergamon Press pic. Allozyme Diversity and lntrogression in the Galapagos Islands Endemic Gossypium datwinii and its Relationship to Continental G. barbadense JONATHAN F. WENDEL* and RICHARD G. PERCYt *Department of Botany, Bessey Hall, Iowa State University, Ames, lA 50011, U.S.A.; tUSDA-ARS, in cooperation with the Arizona Agricultural Experimental Station, Maricopa Agricultural Center, 37860 W. Smith-Enke Rd, Maricopa, AZ 85339, U.S.A. Key Word lndex-Gossypium darwinii; G. barbadense; Malvaceae; cotton; introgression; Galapagos Islands; genetic varia tion; island evolution; allozymes; molecular divergence. Abstract-Gossypium darwiniiWatt is a tetraploid cotton endemic to the Galapagos Islands. Opinion has been divided as to whether or not it deserves recognition at the specific rank, with some considering it a variety of its presumed progenitor, the widely distributed South American species G. barbadense L. A previous hypothesis states that much of the perceived inter gradation between the two taxa arose as a consequence of introgression from G. barbadense following its introduction to the archipelago during the past several hundred years. We performed allozyme analysis on 58 accessions of G. darwinii from six islands, using 17 enzymes collectively encoded by 59 loci. Levels of variation were high for an island endemic, with a mean number of alleles per locus of 1.34 and an average panmictic heterozygosity of 0.062. Principal component analysis revealed clustering of accessions according to their island of origin, and a spatial pattern of island-clusters that approximates geo graphical relationships among islands. Genetic relationships of G. darwiniiwith G. barbadense and G. hirsutum L. were studied using previously generated allozyme data. Significant introgression of G. hirsutum alleles was detected; however morphologi cal considerations support the hypothesis that much of G. darwinii's diversity stems from interspecific gene flow from G. barbadense. Evidence is presented suggesting that the occurrence of G. hirsutum alleles in G. darwinii derives not from direct hybridization, but from a mediated transfer through introduced, G. hirsutum-introgressed G. barbadense. Gossypium darwinii and G. barbadense are nearly fixed for different alleles at four loci and each contains a large number of unique alleles. Notwithstanding the high interspecific Nei's genetic identity (0.949). the allozyme data support geographical and morphological evidence in suggesting that a specific rank for G. darwinii is warranted. Introduction G. darwinii, and its presumed progenitor, is G. Approximately 500 species of vascular plants are barbadense L., one of two commercially import indigenous to the Galapagos Islands, including ant (with G. hirsutum L.) tetraploid (2n=4x=52) about 200 endemics [1-3]. Two of these species of cotton. Gossypium barbadense has an endemics are members of the cotton genus indigenous cultivated range throughout the New ( Gossypium L.). G. k/otzschianum Anderss. World tropics, but its pre-agricultural distribu (reported from five of the islands) and G. darwinii tion appears to be northwestern S. American [5, Watt , (from 13 islands). Phytogeographic 6]. evidence suggests that most of the Galapagos Opinion has been divided as to whether G. flora (approximately 90%) arose from western darwinii deserves recognition at the specific South American ancestors, some 1000 km or rank. Watt, in honoring the Galapagos' most more distant [1-3]. An exception to this generali famous visitor with the commemorative epithet, zation is G. k/otzschianum, one of 13 diploid emphasized several morphological disconti (2n=26) Gossypium species native to the West nuities between the two taxa and their geo ern Hemisphere. Gossypium klotzschianum is graphic isolation [7]. He stated that "At the time unique among the Galapagos endemics in that it at which this species was collected by Darwin [in apparently originated from a northern Mexican 1835 on San Salvador, including the type progenitor [4]. In contrast, the closest relative of specimen] . cotton could hardly have been cultivated anywhere on the Galapagos Islands (Received 25 June 1990) ...", and concluded that "it is a perfectly good 517 518 J. F. WENDEL AND R. G. PERCY species." Later adherents of this view include from throughout the species' range (Table 1, Fig. Harland [8], based on comparative genetic 1). Evidence from previous studies indicated that analyses, and Fryxell [9], in the most recent these 17 enzymes are encoded by a minimum of monograph of the genus. The alternative opinion 59 genetic loci [6]. Allelic variants at 14 of these was promulgated by Hutchinson et al [5], who loci distinguish G. barbadense from G. hirsutum reduced G. darwinii to varietal rank [G. bar [6; Wendel, unpublished data], providing the badense var. darwinii (Watt) J. B. Hutch.]. capability of documenting interspecific introgres Kearney [10] and Valicek [11] concurred, the sion (see Discussion). An initial inspection of the latter author stating that G. darwinii is " ... allelic presence data for the 58 G. darwinii acces decidedly not an independent (separate) sions led to the recognition of seven accessions species." (AS892, CB3087, CB3097, CB3098, WB1207, Hutchinson et al [5] offered several reasons WB1249, PW56) that contained one or more G. for reducing G. darwiniito varietal status, includ hirsutum alleles. These seven accessions were ing its highly variable morphology, the vigorous dropped from the initial data set and are not in and fertile F2 generation obtained from inter cluded in the results presented in Tables 2-5. specific G. barbadense X G. darwinii hybrids, For the