RESUMEN
Spatial patterns of genetic variation can reveal otherwise cryptic evolutionary and landscape processes. In northwestern Costa Rica, an approximately concordant genetic discontinuity occurs among populations of several plant species. We conducted phylogeographic analyses of an epiphytic orchid, Brassavola nodosa, to test for genetic discontinuity and to explore its underlying causes. We genotyped 18 populations with 19 nuclear loci and two non-coding chloroplast sequence regions. We estimated genetic diversity and structure, relative importance of pollen and seed dispersal, and divergence time to understand how genetic diversity was spatially partitioned. Nuclear genetic diversity was high with little differentiation among populations (GSTn = 0.065). In contrast, chloroplast haplotypes were highly structured (GSTc = 0.570) and reveal a discontinuity between northwestern and southeastern populations within Costa Rica. Haplotype differences suggest two formerly isolated lineages that diverged ~10,000-100,000 YBP. Haplotype mixing and greater genetic diversity occur in an intermediate transition zone. Patterns of nuclear and chloroplast data were consistent. Different levels of genetic differentiation for the two genomes reflect the relative effectiveness of biotic versus abiotic dispersers of pollen and seeds, respectively. Isolation of the two lineages likely resulted from the complex environmental and geophysical history of the region. Our results suggest a recent cryptic seed dispersal barrier and/or zone of secondary contact. We hypothesize that powerful northeasterly trade winds hinder movement of wind-borne seeds between the two regions, while the multi-directional dispersal of pollen by strong-flying sphinx moths resulted in lower differentiation of nuclear loci.
Asunto(s)
Variación Genética , Haplotipos , Orchidaceae/genética , Polen/genética , Animales , Costa Rica , Genética de Población , Mariposas Nocturnas , Orchidaceae/crecimiento & desarrollo , Filogeografía , Polinización , Dispersión de SemillasRESUMEN
PREMISE OF THE STUDY: Flowering initiation, duration and magnitude, and degree of flowering synchrony within a population can affect the reproductive fitness of individuals. We examined the flowering phenology within a population of the tropical dry forest Guanacaste tree (Enterolobium cyclocarpum) to gauge the impact of phenological variation among trees on fruit production and progeny vigor. METHODS: We monitored the flowering phenology of 93 trees weekly during 2005, 2006, and 2007, using a scale based on the percentage of the crown with open flowers. We also monitored fruit production for each tree in 2005, 2006, 2007, and 2008. Finally, we evaluated the relationship between phenological variation and progeny performance. KEY RESULTS: Ten measures of flowering phenology and synchrony among flowering trees, based on the number of weeks when anthesis of the crown exceeded 50%, were used to develop four phenological profiles. These profiles were correlated with significant differences in fruit production and progeny vigor. Trees with flowers in >50% of their crown for at least 2 weeks produced more fruits and more vigorous progeny than trees with other profiles. Trees also tended to produce the same phenological profile among years than predicted by chance. CONCLUSIONS: Guanacaste trees vary significantly in the initiation of anthesis, duration and magnitude of flowering, and degree of synchrony among trees. Trees also tend to maintain the same flowering profile among years. Finally, the flowering behavior of E. cyclocarpum leads to significant differences in fruit and seed production, germination, and early progeny growth.
Asunto(s)
Fabaceae/fisiología , Flores/fisiología , Árboles/fisiología , Frutas/crecimiento & desarrollo , Germinación , Plantones/crecimiento & desarrollo , Factores de TiempoRESUMEN
Populations of many species are isolated within narrow elevation bands of Neotropical mountain habitat, and how well dispersal maintains genetic connectivity is unknown. We asked whether genetic structure of an epiphytic orchid, Epidendrum firmum, corresponds to gaps between Costa Rican mountain ranges, and how these gaps influence pollen and seed flow. We predicted that significant genetic structure exists among mountain ranges due to different colonization histories and limited gene flow. Furthermore, we predicted that pollen movement contributes more to gene flow than seeds because seeds are released into strong winds perpendicular to the narrow northwest-southeast species distribution, while the likely pollinators are strong fliers. Individuals from 12 populations and three mountain ranges were genotyped with nuclear microsatellites (nDNA) and chloroplast sequences (cpDNA). Genetic diversity was high for both markers, while nDNA genetic structure was low (FSTn = 0.020) and cpDNA structure was moderate (FSTc = 0.443). Significant cpDNA barriers occurred within and among mountain ranges, but nDNA barriers were not significant after accounting for geographic distance. Consistent with these contrasting patterns of genetic structure, pollen contributes substantially more to gene flow among populations than seed (mp /ms = 46). Pollinators mediated extensive gene flow, eroding nDNA colonization footprints, while seed flow was comparatively limited, possibly due to directional prevailing winds across linearly distributed populations. Dispersal traits alone may not accurately inform predictions about gene flow or genetic structure, supporting the need for research into the potentially crucial role of pollinators and landscape context in gene flow among isolated populations.
Asunto(s)
Flujo Génico , Variación Genética , Orchidaceae/genética , Polinización , Dispersión de Semillas , Núcleo Celular/genética , Costa Rica , ADN de Cloroplastos/genética , Ecosistema , Genética de Población , Genotipo , Geografía , Repeticiones de Microsatélite , Datos de Secuencia Molecular , Polen/genética , VientoRESUMEN
Symbiotic interactions are common in nature. In dynamic or degraded environments, the ability to associate with multiple partners (i.e. broad specificity) may enable species to persist through fluctuations in the availability of any particular partner. Understanding how species interactions vary across landscapes is necessary to anticipate direct and indirect consequences of environmental degradation on species conservation. We asked whether mycorrhizal symbiosis by populations of a rare epiphytic orchid (Epidendrum firmum) is related to geographic or environmental heterogeneity. The latter would suggest that interactions are governed by environmental conditions rather than historic isolation of populations and/or mycorrhizal fungi. We used DNA-based methods to identify mycorrhizal fungi from eleven E. firmum populations in Costa Rica. We used molecular and phylogenetic analyses to compare associations. Epidendrum firmum exhibited broad specificity, associating with diverse mycorrhizal fungi, including six Tulasnellaceae molecular operational taxonomic units (MOTUs), five Sebacinales MOTUs and others. Notably, diverse mycorrhizal symbioses formed in disturbed pasture and roadside habitats. Mycorrhizal fungi exhibited significant similarity within populations (spatial and phylogenetic autocorrelation) and significant differences among populations (phylogenetic community dissimilarity). However, mycorrhizal symbioses were not significantly associated with biogeographic or environmental features. Such unexpected heterogeneity among populations may result from complex combinations of fine-scale environmental factors and macro-evolutionary patterns of change in mycorrhizal specificity. Thus, E. firmum exhibits broad specificity and the potential for opportunistic associations with diverse fungi. We suggest that these characteristics could confer symbiotic assurance when mycorrhizal fungi are stochastically available, which may be crucial in dynamic or disturbed habitats such as tropical forest canopies.
Asunto(s)
Micorrizas/clasificación , Orchidaceae/microbiología , Filogenia , Simbiosis , Ascomicetos/clasificación , Ascomicetos/genética , Basidiomycota/clasificación , Basidiomycota/genética , Biodiversidad , Costa Rica , Código de Barras del ADN Taxonómico , Datos de Secuencia Molecular , Micorrizas/genética , Análisis de Secuencia de ADN , Especificidad de la EspecieRESUMEN
Colonization of vacant habitat is a fundamental ecological process that affects the ability of species to persist and undergo range modifications in continually shifting landscapes. Thus, understanding factors that affect and limit colonization has important ecological and conservation implications. Epiphytic orchids are increasingly threatened by various factors, including anthropogenic habitat disturbance. As cleared areas (e.g. pastures) are recolonized by suitable host trees, the establishment and genetic composition of epiphytic orchid populations are likely a function of their colonization patterns. We used genetic analyses to infer the prevailing colonization pattern of the epiphytic orchid, Brassavola nodosa. Samples from three populations (i.e. individuals within a tree) from each of five pastures in the dry forest of Costa Rica were genotyped with neutral nuclear and chloroplast markers. Spatial autocorrelation and hierarchical genetic structure analyses were used to assess the relatedness of individuals within populations, among populations within pastures and among populations in different pastures. The results showed significant relatedness within populations (mean r=0.166) and significant but lower relatedness among populations within a pasture (mean r=0.058). Our data suggest that colonization of available habitats is by few individuals with subsequent population expansion resulting from in situ reproduction, and that individuals within a tree are not a random sample of the regional seed pool. Furthermore, populations within a pasture were likely colonized by seeds produced by founders of a neighbouring population within that pasture. These results have important ramifications for understanding conservation measures needed for this species and other epiphytic orchids.
Asunto(s)
Ecología , Genética de Población , Orchidaceae , Costa Rica , Ecosistema , Genotipo , Humanos , SemillasRESUMEN
PREMISE OF THE STUDY: Ten microsatellite loci were isolated and characterized for the neotropical epiphytic orchid Epidendrum firmum to examine levels of genetic diversity and genetic structure at multiple spatial scales. METHODS AND RESULTS: We screened loci in 12-25 individuals from each of two populations in Costa Rica and identified 10 polymorphic loci. The number of alleles per locus ranged from one to 15 while observed heterozygosity for polymorphic loci ranged from 0.360 to 0.960. CONCLUSIONS: Primers for these informative genetic markers will be useful for quantifying genetic diversity, spatial genetic structure, and gene flow in E. firmum.
Asunto(s)
Cartilla de ADN/genética , Variación Genética , Repeticiones de Microsatélite/genética , Orchidaceae/genética , Alelos , Costa Rica , ADN de Plantas/química , ADN de Plantas/genética , Genotipo , Datos de Secuencia Molecular , Reacción en Cadena de la Polimerasa , Polimorfismo Genético , Análisis de Secuencia de ADNRESUMEN
We examined spatial genetic structure (SGS) in Enterolobium cyclocarpum (the Guanacaste tree), a dominant tree of Central American dry forests in 4 sites in Guanacaste Province, Costa Rica. In disturbed dry forest sites (e.g., pastures), E. cyclocarpum is primarily dispersed by cattle and horses, whose movements are restricted by pasture boundaries. The study sites varied in tree densities and disturbance. Allozyme analyses of adult trees demonstrated significant levels of SGS in 3 of 4 sites. SGS was primarily due to clusters of young adults located along seasonal streams, rocky areas, and in abandoned pastures. SGS was highest in the first distance class in the least disturbed population, which also had the lowest density of large adults. Low, but significant SGS characterized the site with the highest number of large adults located in individual pastures. The semiurban site, had no clusters of young adults and, probably as a result, failed to exhibit SGS. Our results demonstrate that disturbance can strongly influence SGS patterns and are consistent with a landscape model in which the location of potential recruitment sites, restricted seed disperser movements, and the number and location of maternal individuals dictate the level and pattern of SGS.
Asunto(s)
Ambiente , Fabaceae/genética , Genética de Población , Costa Rica , Demografía , Ecosistema , Fabaceae/fisiología , Flujo Génico/fisiología , Especiación Genética , Variación Genética/fisiología , ÁrbolesRESUMEN
Insights into processes that lead to the distribution of genetic variation within plant species require recognition of the importance of both pollen and seed movement. Here we investigate the contributions of pollen and seed movement to overall gene flow in the Central American epiphytic orchid, Laelia rubescens. Genetic diversity and structure were examined at multiple spatial scales in the tropical dry forest of Costa Rica using nuclear (allozymes) and chloroplast restriction fragment length polymorphism (RFLP) markers, which were found to be diverse (allozymes, P = 73.3%; HE = 0.174; cpDNA, HE = 0.741). Nuclear genetic structure (FSTn) was low at every spatial scale (0.005-0.091). Chloroplast markers displayed more structure (0.073-0.254) but relatively similar patterns. Neither genome displayed significant isolation-by-distance. Pollen and seed dispersal rates did not differ significantly from one another (mp/ms = 1.40) at the broadest geographical scale, among sites throughout Costa Rica. However, relative contributions of pollen and seeds to gene flow were scale-dependent, with different mechanisms determining the dominant mode of gene flow at different spatial scales. Much seed dispersal is highly localized within the maternal population, while some seeds enter the air column and are dispersed over considerable distances. At the intermediate scale (10s to 100s of metres) pollinators are responsible for substantial pollen flow. This species appears capable of distributing its genes across the anthropogenically altered landscape that now characterizes its Costa Rican dry forest habitat.
Asunto(s)
Variación Genética , Genética de Población , Orchidaceae/genética , Polen/fisiología , Semillas/fisiología , Costa Rica , ADN de Cloroplastos/genética , Geografía , Isoenzimas , Desequilibrio de Ligamiento , Orchidaceae/fisiología , Polimorfismo de Longitud del Fragmento de Restricción , Dinámica PoblacionalRESUMEN
Epiphytic plants occupy three-dimensional space, which allows more individuals to be closely clustered spatially than is possible for populations occupying two dimensions. The unique characteristics of epiphytes can act in concert to influence the fine-scale genetic structure of their populations which can, in turn, influence mating patterns and other population phenomena. Three large populations of Laelia rubescens (Orchidaceae) in the Costa Rican seasonal dry forest were sampled at two levels of intensity to determine: (i) whether individual clusters contain more than one genotype, and (ii) the spatial distribution and fine-scale genetic structure of genotypes within populations. Samples were assayed for their multilocus allozyme genotypes and spatial autocorrelation analyses were performed. High levels of genetic diversity, high genotypic diversity and low among-population variation were found. In the larger clusters, multiple genets per cluster were common with discrete clusters containing up to nine genotypes. Spatial autocorrelation analyses indicated significant positive genetic structure at distances of = 45 cm. This result is likely due to the formation of discrete clusters by vegetative reproduction, as well as the establishment of sexually derived progeny within and near maternal clusters.