Genetic structure of coexisting wild and managed agave populations: implications for the evolution of plants under domestication.

Domestication is a continuous evolutionary process guided by humans. This process leads to divergence in characteristics such as behaviour, morphology or genetics, between wild and managed populations. Agaves have been important resources for Mesoamerican peoples since prehistory. Some species are domesticated and others vary in degree of domestication. Agave inaequidens Koch is used in central Mexico to produce mescal, and a management gradient from gathered wild and silvicultural populations, as well as cultivated plantations, has been documented. Significant morphological differences were reported among wild and managed populations, and a high phenotypic variation in cultivated populations composed of plants from different populations. We evaluated levels of genetic diversity and structure associated with management, hypothesizing that high morphological variation would be accompanied by high genetic diversity in populations with high gene flow and low genetic structure among managed and unmanaged populations. Wild, silvicultural and cultivated populations were studied, collecting tissue of 19-30 plants per population. Through 10 nuclear microsatellite loci, we compared population genetic parameters. We analysed partition of variation associated with management categories to estimate gene flow among populations. Agave inaequidens exhibits high levels of genetic diversity (He = 0.707) and moderate genetic structure (FST = 0.112). No differences were found in levels of genetic diversity among wild (He = 0.704), silviculturally managed (He = 0.733) and cultivated (He = 0.698) populations. Bayesian analysis indicated that five genetic clusters best fit the data, with genetic groups corresponding to habitats where populations grow rather than to management. Migration rates ranged from zero between two populations to markedly high among others (M = 0.73-35.25). Natural mechanisms of gene flow and the dynamic management of agave propagules among populations favour gene flow and the maintenance of high levels of variation within all populations. The slight differentiation associated with management indicates that domestication is in an incipient stage.

Morphological variation, management and domestication of 'maguey alto' (Agave inaequidens) and 'maguey manso' (A. hookeri) in Michoacán, México.

BACKGROUND: Agave inaequidens and A. hookeri are anciently used species for producing the fermented beverage 'pulque', food and fiber in central Mexico. A. inaequidens is wild and cultivated and A. hookeri only cultivated, A. inaequidens being its putative wild relative. We analysed purposes and mechanisms of artificial selection and phenotypic divergences between wild and managed populations of A. inaequidens and between them and A. hookeri, hypothesizing phenotypic divergence between wild and domesticated populations of A. inaequidens in characters associated to domestication, and that A. hookeri would be phenotypically similar to cultivated A. inaequidens. METHODS: We studied five wild and five cultivated populations of A. inaequidens, and three cultivated populations of A. hookeri. We interviewed agave managers documenting mechanisms of artificial selection, and measured 25 morphological characters. Morphological similarity and differentiation among plants and populations were analysed through multivariate methods and ANOVAs. RESULTS: People recognized 2-8 variants of A. inaequidens; for cultivation they select young plants collected in wild areas recognized as producing the best quality mescal agaves. Also, they collect seeds of the largest and most vigorous plants, sowing seeds in plant beds and then transplanting the most vigorous plantlets into plantations. Multivariate methods classified separately the wild and cultivated populations of A. inaequidens and these from A. hookeri, mainly because of characters related with plant and teeth size. The cultivated plants of A. inaequidens are significantly bigger with larger teeth than wild plants. A. hookeri are also significatly bigger plants with larger leaves but lower teeth density and size than A. inaequidens. Some cultivated plants of A. inaequidens were classified as A. hookeri, and nearly 10% of A. hookeri as cultivated A. inaequidens. Wild and cultivated populations of A. inaequidens differed in 13 characters, whereas A. hookeri differed in 23 characters with wild populations and only in 6 characters with cultivated populations of A. inaequidens. CONCLUSIONS: Divergence between wild and cultivated populations of A. inaequidens reflect artificial selection. A. hookeri is similar to the cultivated A. inaequidens, which supports the hypothesis that A. hookeri could be the extreme of a domestication gradient of a species complex.

[Interpopulation reproductive synchrony of Agave cocui (Agavaceae) in Venezuela]. / Sincronía reproductiva interpoblacional de Agave cocui (Agavaceae) en Venezuela.

Agave cocui (Agavaceae) is a species with broad distribution in arid and semiarid areas of Venezuela and Colombia. Despite of its ecological importance as a source of food for wildlife, and its economic value for production of a spirit drink, studies on the reproductive ecology of the species are relatively rare. In this study, we conducted a one-year evaluation of the flowering and fruiting phenology of A. cocui in the eight representative localities of the species' distribution in Venezuela. Within each study site, we chose an area with a minimum of 50 reproductive individuals and followed their reproductive phenophases with the help of binoculars, using six qualitative cathegories (emerging reproductive stalk, flowers, inmature fruits, mature fruits, bulbils and dry stalk) every two months. Emergence of the reproductive stalk in most of the examined populations began in September (rainy season), although this event delayed two months in a few populations. We detected significant negative correlations between precipitation and the percentage of flowering occurrence in four of the eight populations. Floral resources are available for flower visitors during approximately five months of the year (January-May). In most populations production of flowers initiated in January (dry season), and for Western Venezuela and Andean regions, the flowering main peak occurred in January. Localities from the Central and Eastern Coast exhibited the flowering peak in March, showing a delay of approximately two months with respect to other populations. Beginning of fruit set varied among localities from January to May; however, peak production of mature fruits concentrated in May, and fruit occurrence varied broadly between 5.2 and 85%. Bulbil production was detected in all populations and varied greatly among them (maximum percentage per population: 26.19-92.10%). High flowering synchronicity (Phenophase Overlapping Index: 0.756 and 0.999) was observed among all populations monitored in Western Venezuela, including the Andean localities. This condition might facilitate the existence of a nectar corridor from the Western Coast and nearby islands, to the Andean arid patches, which could be potentially used by nectar-feeding bats and birds dependent on agave flowers during part of the year.

Sincronía reproductiva interpoblacional de Agave cocui (Agavaceae) en Venezuela

Interpopulation reproductive synchrony of Agave cocui (Agavaceae) in Venezuela. Agave cocui (Agavaceae) is a species with broad distribution in arid and semiarid areas of Venezuela and Colombia. Despite of its ecological importance as a source of food for wildlife, and its economic value for production of a spirit drink, studies on the reproductive ecology of the species are relatively rare. In this study, we conducted a oneyear evaluation of the flowering and fruiting phenology of A. cocui in the eight representative localities of the species’ distribution in Venezuela. Within each study site, we chose an area with a minimum of 50 reproductive individuals and followed their reproductive phenophases with the help of binoculars, using six qualitative cathegories (emerging reproductive stalk, flowers, inmature fruits, mature fruits, bulbils and dry stalk) every two months. Emergence of the reproductive stalk in most of the examined populations began in September (rainy season), although this event delayed two months in a few populations. We detected significant negative correlations between precipitation and the percentage of flowering occurrence in four of the eight populations. Floral resources are available for flower visitors during approximately five months of the year (January-May). In most populations production of flowers initiated in January (dry season), and for Western Venezuela and Andean regions, the flowering main peak occurred in January. Localities from the Central and Eastern Coast exhibited the flowering peak in March, showing a delay of approximately two months with respect to other populations. Beginning of fruit set varied among localities from January to May; however, peak production of mature fruits concentrated in May, and fruit occurrence varied broadly between 5.2 and 85%. Bulbil production was detected in all populations and varied greatly among them (maximum percentage per population: 26.19-92.10%). High flowering synchronicity (Phenophase Overlapping Index: 0.756 and 0.999) was observed among all populations monitored in Western Venezuela, including the Andean localities. This condition might facilitate the existence of a nectar corridor from the Western Coast and nearby islands, to the Andean arid patches, which could be potentially used by nectar-feeding bats and birds dependent on agave flowers during part of the year. Rev. Biol. Trop. 59 (3): 1359-1370. Epub 2011 September 01.

cocui (Agavaceae) es una especie de amplia distribución en zonas áridas y semiáridas de Venezuela y Colombia. A pesar de su importancia ecológica como fuente de recursos para la fauna silvestre y de su potencial valor económico como materia prima para la producción del licor “Cocuy de Penca”, los estudios sobre su ecología reproductiva son muy escasos. En este estudio se evaluó la fenología de floración y fructificación de A. cocui en ocho localidades de Venezuela. La fase reproductiva se inició al finalizar el periodo de lluvias. Las flores se producen durante cinco meses. Para la Región Noroccidental del país y los Andes los máximos de floración ocurrieron en enero, mientras que las localidades en la costa central y oriental exhibieron un desface de cerca de dos meses, a principios de marzo. La mayor sincronía floral ocurrió entre todas las poblaciones censadas en el occidente del país, incluyendo las localidades andinas. Dicha sincronía reproductiva potenciaría la conformación de un corredor de néctar para aves y murciélagos, que se extendería desde el Norte de Falcón, en la Costa Occidental, hasta los bolsones andinos, en el Suroeste del país.

Population genetics of Agave cocui: evidence for low genetic diversity at the southern geographic limit of genus Agave.

The Agave genus embraces many species with outstanding ecological and economic importance in the arid regions of the Americas. Even though this genus covers a broad geographic distribution, our knowledge on the population genetics of species is concentrated in taxa located in North America. Recently, it has been demonstrated that plant domestication decreases levels of genetic diversity in managed populations and increases population structure with respect to wild populations. We examined levels of allozyme diversity (N = 17 loci) and population structure of Agave cocui, the species at the southern limit of distribution of the genus. We sampled 7 wild populations (N = 30-35 individuals per population) representative of the geographic distribution of the species in Venezuela. Among the agaves studied, A. cocui has some of the lowest estimates of genetic diversity (H(e)[species] = 0.059, H(e)[population] = 0.054) reported until present. We propose that this condition is probably linked to the recent origin of this species in arid and semiarid regions of Colombia and Venezuela, probably through one or a few founder events. The lowest estimates of genetic diversity were associated with small populations in very restricted arid patches; but also with overexploitation of rosettes for production of fermented drinks and fibers. Santa Cruz de Pecaya, one of the 2 centers of economic use of agaves in northwestern Venezuela presented one of the lowest values of genetic variability, a sign suggesting that human impact represents a significant threat to the available genetic pool that this species possesses in the region.