Phylogenomic Analysis of the Explosive Adaptive Radiation of the Espeletia Complex (Asteraceae) in the Tropical Andes.

The subtribe Espeletiinae (Asteraceae), endemic to the high-elevations in the Northern Andes, exhibits an exceptional diversity of species, growth-forms, and reproductive strategies. This complex of 140 species includes large trees, dichotomous trees, shrubs and the extraordinary giant caulescent rosettes, considered as a classic example of adaptation in tropical high-elevation ecosystems. The subtribe has also long been recognized as a prominent case of adaptive radiation, but the understanding of its evolution has been hampered by a lack of phylogenetic resolution. Herein, we produce the first fully resolved phylogeny of all morphological groups of Espeletiinae, using whole plastomes and about a million nuclear nucleotides obtained with an original de novo assembly procedure without reference genome, and analyzed with traditional and coalescent-based approaches that consider the possible impact of incomplete lineage sorting and hybridization on phylogenetic inference. We show that the diversification of Espeletiinae started from a rosette ancestor about 2.3 Ma, after the final uplift of the Northern Andes. This was followed by two independent radiations in the Colombian and Venezuelan Andes, with a few trans-cordilleran dispersal events among low-elevation tree lineages but none among high-elevation rosettes. We demonstrate complex scenarios of morphological change in Espeletiinae, usually implying the convergent evolution of growth-forms with frequent loss/gains of various traits. For instance, caulescent rosettes evolved independently in both countries, likely as convergent adaptations to life in tropical high-elevation habitats. Tree growth-forms evolved independently three times from the repeated colonization of lower elevations by high-elevation rosette ancestors. The rate of morphological diversification increased during the early phase of the radiation, after which it decreased steadily towards the present. On the other hand, the rate of species diversification in the best-sampled Venezuelan radiation was on average very high (3.1 spp/My), with significant rate variation among growth-forms (much higher in polycarpic caulescent rosettes). Our results point out a scenario where both adaptive morphological evolution and geographical isolation due to Pleistocene climatic oscillations triggered an exceptionally rapid radiation for a continental plant group.

Plant diversity patterns in neotropical dry forests and their conservation implications.

Seasonally dry tropical forests are distributed across Latin America and the Caribbean and are highly threatened, with less than 10% of their original extent remaining in many countries. Using 835 inventories covering 4660 species of woody plants, we show marked floristic turnover among inventories and regions, which may be higher than in other neotropical biomes, such as savanna. Such high floristic turnover indicates that numerous conservation areas across many countries will be needed to protect the full diversity of tropical dry forests. Our results provide a scientific framework within which national decision-makers can contextualize the floristic significance of their dry forest at a regional and continental scale.

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.

Phyllostomid bat occurrence in successional stages of neotropical dry forests.

Tropical dry forests (TDFs) are highly endangered tropical ecosystems being replaced by a complex mosaic of patches of different successional stages, agricultural fields and pasturelands. In this context, it is urgent to understand how taxa playing critical ecosystem roles respond to habitat modification. Because Phyllostomid bats provide important ecosystem services (e.g. facilitate gene flow among plant populations and promote forest regeneration), in this study we aimed to identify potential patterns on their response to TDF transformation in sites representing four different successional stages (initial, early, intermediate and late) in three Neotropical regions: México, Venezuela and Brazil. We evaluated bat occurrence at the species, ensemble (abundance) and assemblage level (species richness and composition, guild composition). We also evaluated how bat occurrence was modulated by the marked seasonality of TDFs. In general, we found high seasonal and regional specificities in phyllostomid occurrence, driven by specificities at species and guild levels. For example, highest frugivore abundance occurred in the early stage of the moistest TDF, while highest nectarivore abundance occurred in the same stage of the driest TDF. The high regional specificity of phyllostomid responses could arise from: (1) the distinctive environmental conditions of each region, (2) the specific behavior and ecological requirements of the regional bat species, (3) the composition, structure and phenological patterns of plant assemblages in the different stages, and (4) the regional landscape composition and configuration. We conclude that, in tropical seasonal environments, it is imperative to perform long-term studies considering seasonal variations in environmental conditions and plant phenology, as well as the role of landscape attributes. This approach will allow us to identify potential patterns in bat responses to habitat modification, which constitute an invaluable tool for not only bat biodiversity conservation but also for the conservation of the key ecological processes they provide.

Trypanosoma evansi kDNA minicircle found in the Venezuelan nectar-feeding bat Leptonycteris curasoae (Glossophaginae), supports the hypothesis of multiple origins of that parasite in South America.

Trypanosoma evansi is a mammal generalist protozoon which causes negative effects on health and productivity in bovine and equine herds in South America, Europe, Asia and Africa. By molecular methods, we screened the presence of that parasite together with other trypanosome species in 105 bats of 10 species collected in arid zones of northern Venezuela. The first molecular approach was fluorescent fragment length barcoding (FFLB), which relies on amplification of relative small regions of rRNA genes (four loci) and fluorescence detection. By FFLB, 17 samples showed patterns of possible trypanosomatid infections. These samples were used to test presence of trypanosomes by PCR using the following DNA markers: V7-V8 SSU rRNA, gGAPDH and kDNA minicircle regions. Only in one individual of the nectar-feeding bat, Leptonycteris curasoae, we were able to amplify 1000bp of the trypanosome kDNA minicircle. That PCR product was sequenced and the parasite species was determined by NCBI-BLAST and phylogenetic analysis. Both analyses showed that the minicircle sequence corresponds to Trypanosoma evansi. The phylogenetic analysis of the sequence obtained in this study clustered with a T. evansi sequence obtained in a Venezuelan capybara, Hydrochoerus hydrochaeris, and distant of others two T. evansi sequences obtained in a Colombian capybara and horse. This result supports the hypothesis of multiple origins of T. evansi in South America.

[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.

Seasonal sebaceous patch in the nectar-feeding bats Leptonycteris curasoae and L. yerbabuenae (Phyllostomidae: Glossophaginae): phenological, histological, and preliminary chemical characterization.

Many species of bats secrete a wide variety of substances, frequently associated with olfactory communication. We characterized a seasonal phenomenon of dorsal sebaceous secretion in the Curaçaoan long-nosed bat, Leptonycteris curasoae, in Venezuela, and the lesser long-nosed bat, Leptonycteris yerbabuenae, in Mexico. The phenology of the sebaceous patch was determined, a histological analysis of the affected area was conducted using specimens of L. curasoae from Venezuela, and finally, a preliminary chemical characterization of the substance secreted was performed combining histochemical techniques with gas chromatography and mass spectrometry analyses. The sebaceous patch was detected exclusively in male adult specimens. Individuals presenting it had a variable area of fur covered with a fatty and odoriferous substance at the level of the interscapular zone. Occurrence of the sebaceous patch was cyclical and coincided with the mating season in Venezuela and Mexico. The following histological changes associated with occurrence of the patch were observed: increase of epidermis thickness and decrease of dermis and hypodermis thicknesses, increase in density of sebaceous glands, increase of percentage of skin covered by sebaceous glands, increase of size of sebaceous glands previous to secretion followed, and increase of the sebum volume within sebaceous glands previous to secretion. Several compounds tentatively identified as fatty acids, cholestanes and cholesterol were present in the sebaceous secretion. Based on the evidence obtained, we hypothesize that the sebaceous patch could be involved in olfactory communication, possibly related to mating behavior in these bats.

Reproductive biology and mating system estimates of two Andean melocacti, Melocactus schatzlii and M. andinus (Cactaceae).

BACKGROUND AND AIMS: The genus Melocactus comprises 36 species of globose cacti with the most derived traits in the Cereeae tribe. It is the proper study system to examine what are the most derived reproductive strategies within that tribe. This study aims to characterize the reproductive biology and to estimate the mating system parameters of two Andean melocacti, Melocactus schatzlii and M. andinus. METHODS: The reproductive attributes of the two species were described, including floral morphology, anthesis patterns, floral rewards, floral visitors and visitation patterns. Levels of self-compatibility and autonomous self-pollination were estimated by hand-pollination experiments. Mating system estimates were obtained by conducting progeny array analyses using isozymes. KEY RESULTS: The flowers of the two species present the typical hummingbird-pollination syndrome. Despite their morphological resemblance, the two species differ in flower size, pollen and ovule production and anthesis pattern. Their main pollinator agents are hummingbirds, four species in M. schatzlii and one species in M. andinus. Both cacti are self-compatible and capable of self-pollination without the aid of pollen vectors. Population-level outcrossing rate was higher for M. schatzlii (t(m)=0.9) than for M. andinus (t(m)=0.4). At the family level, outcrossing rates for most mothers of M. schatzlii were higher (t(m)>0.8) than for M. andinus (t(m)<0.5). CONCLUSIONS: Although the two cacti are capable of selfing, M. schatzlii is a predominantly outcrossing species, while M. andinus behaves as a mixed-mating cactus. Hummingbirds are the only pollinators responsible for outcrossing and gene flow events in these species. In their absence, both melocacti set seeds by selfing. Based on its low population size, restricted distribution in Venezuela, low rates of floral visits, and high levels of inbreeding, M. andinus is considered to be an endangered species deserving further study to define its conservation status.

[Cardon dato (Stenocereus griseus, Cactaceae) fruit pulp as raw material for marmalade production]. / Pulpa del fruto del cardón dato (Stenocereus griseus, Cactaceae) como materia prima para la elaboración de mermelada.

This study aimed to examine the possibility of producing marmalades from the fruit pulp of the red and white varieties of card6n dato (Stenocereus griseus). We developed the formulations of the marmalades and evaluated their stability during three months of storage at room temperature. As first step, we characterized the fruits of the two color varieties, observing that despite the considerable difference between both color varieties, there were no significant differences in average weight, dimensions and proportion of pulp, skin and seeds. The pulp of the two color types had high pH (5.2) values and low contents of reductor sugars (3.59 g/ 100 g white variety 2.23 g/ 100 g red variety), non reductor sugars (0.75 g/ 100 g white variety and 2.03 g/ 100 g red variety), pectin (0.14 g/ 100 g white variety and 0.23 g/ 100 g red variety) and acids (7.67 g/ 100 g white variety and 0.15 g/ 100 g red variety). It is needed to include sugar, pectin, and citric acid in the marmalade formula. The first marmalades produced were gummy, a problem that we solved adding the acid from the beginning of the making process. During the three months of storage, the marmalades had good acceptance by the evaluators, this despite slight fluctuations observed in pH, solid contents, and acidity.

Pulpa del fruto del cardón dato (Stenocereus griseus, Cactaceae) como materia prima para la elaboración de mermelada / Cardon dato (Stenocereus griseus, Cactaceae) fruit pulp as raw material for marmalade production

El presente trabajo de investigación se evaluó la posibilidad de elaborar mermeladas a partir de la pulpa de los frutos de cardón dato (Stenocereus griseus) de las variedades Blanca y Roja. Para ello, se propuso desarrollar las formulaciones de dichas mermeladas y estudiar su estabilidad durante tres meses de almacenamiento a temperatura ambiente. Como primer paso se caracterizaron los frutos de ambas variedades, observándose que a pesar de la diferencia notable de color entre ambos, no hay diferencias estadísticamente significativas en su peso promedio, dimensiones y proporciones de pulpa, piel y semillas. Por otro lado, ambas pulpas presentaron valores altos de pH (5,2 en ambas variedades) y contenidos bajos de azúcares reductores (3,59 g/100g variedad Blanca y 2,23 g/100g variedad Roja), azúcares no reductores (0,75 g/100g variedad Blanca y 2,03 g/100g variedad Roja), pectina (0,14 g/100g variedad Blanca y 0,23 g/100g variedad Roja) y ácidos (7,67 g/100g variedad Blanca y 0,15 g/100g variedad Roja), debido a lo cual fue necesario añadir azúcar, pectina y ácido cítrico a la formulación de la mermelada. La primeras mermeladas elaboradas presentaron consistencia gomosa, lo cual se solucionó añadiendo el ácido desde el principio del proceso de elaboración en lugar de al final. Durante los tres meses de almacenamiento, las mermeladas tuvieron buena aceptación por parte de los panelistas, a pesar de ligeras fluctuaciones observadas durante el almacenamiento en el pH y contenidos de sólidos solubles y acidez total titulable.

This study aimed to examine the possibility of producing marmalades from the fruit pulp of the red and white varieties of cardón dato (Stenocereus griseus). We developed the formulations of the marmalades and evaluated their stability during three months of storage at room temperature. As first step, we characterized the fruits of the two color varieties, observing that despite the considerable difference between both color varieties, there were no significant differences in average weight, dimensions and proportion of pulp, skin and seeds. The pulp of the two color types had high pH (5,2) values and low contents of reductor sugars (3,59 g/100g white variety 2,23 g/100g red variety), non reductor sugars (0,75 g/100g white variety and 2,03 g/100g red variety), pectin (0,14 g/100g white variety and 0,23 g/100g red variety) and acids (7,67 g/100g white variety and 0,15 g/100g red variety). It is needed to include sugar, pectin, and citric acid in the marmalade formula. The first marmalades produced were gummy, a problem that we solved adding the acid from the beginning of the making process. During the three months of storage, the marmalades had good acceptance by the evaluators, this despite slight fluctuations observed in pH, solid contents, and acidity.

Genetic population structure and mobility of two nectar-feeding bats from Venezuelan deserts: inferences from mitochondrial DNA.

Glossophaga longirostris and Leptonycteris curasoae are nectar-feeding bats associated with arid zones in northern South America. Despite their close phylogenetic relationship, sympatric condition and niche similarities, morphological and ecological evidence suggest that these species differ in dispersal capabilities. Using mitochondrial DNA, we tested the hypothesis that these species exhibit different levels of population structure that are congruent with their particular movement capabilities. We sequenced a section of the control region of mtDNA for 41 G. longirostris and 42 L. curasoae from 11 zones in Venezuela. Population subdivision in G. longirostris (FST = 0.725) was considerably higher than in L. curasoae (FST = 0.167). L. curasoae individuals shared haplotypes at greater distances (812 km) than G. longirostris (592 km). Our results offer preliminary evidence for one of two possible scenarios, either greater mobility in L. curasoae or a higher degree of female philopatry in G. longirostris.

Population genetic structure of Venezuelan chiropterophilous columnar cacti (Cactaceae).

We conducted allozyme surveys of three Venezuelan self-incompatible chiropterophilous columnar cacti: two diploid species, Stenocereus griseus and Cereus repandus, and one tetraploid, Pilosocereus lanuginosus. The three cacti are pollinated by bats, and both bats and birds disperse seeds. Population sampling comprised two spatial scales: all Venezuelan arid zones (macrogeographic) and two arid regions in northwestern Venezuela (regional). Ten to 15 populations and 17-23 loci were analyzed per species. Estimates of genetic diversity were compared with those of other allozyme surveys in the Cactaceae to examine how bat-mediated gene dispersal affects the population genetic attributes of the three cacti. Genetic diversity was high for both diploid (P(s) = 94.1-100, P(p) = 56.7-72.3, H(s) = 0.182-0.242, H(p) = 0.161-0.205) and tetraploid (P(s) = 93.1, P(p) = 76.1, H(s) = 0.274, H(p) = 0.253) species. Within-population heterozygote deficit was detected in the three cacti at macrogeographic (F(IS) = 0.145-0.182) and regional (F(IS) = 0.057-0.174) levels. Low genetic differentiation was detected at both macrogeographic (G(ST) = 0.043-0.126) and regional (G(ST) = 0.009-0.061) levels for the three species, suggesting substantial gene flow among populations. Gene exchange among populations seems to be regulated by distance among populations. Our results support the hypothesis that bat-mediated gene dispersal confers high levels of genetic exchange among populations of the three columnar cacti, a process that enhances levels of genetic diversity within their populations.