Jorge A. Lobo

Genetic consequences of habitat fragmentation in plant populations: susceptible signals in plant traits and methodological approaches

Conservation of genetic diversity, one of the three main forms of biodiversity, is a fundamental concern in conservation biology as it provides the raw material for evolutionary change and thus the potential to adapt to changing environments. By means of meta‐analyses, we tested the generality of the hypotheses that habitat fragmentation affects genetic diversity of plant populations and that certain life history and ecological traits of plants can determine differential susceptibility to genetic erosion in fragmented habitats. Additionally, we assessed whether certain methodological approaches used by authors influence the ability to detect fragmentation effects on plant genetic diversity. We found overall large and negative effects of fragmentation on genetic diversity and outcrossing rates but no effects on inbreeding coefficients. Significant increases in inbreeding coefficient in fragmented habitats were only observed in studies analyzing progenies. The mating system and the rarity status of plants explained the highest proportion of variation in the effect sizes among species. The age of the fragment was also decisive in explaining variability among effect sizes: the larger the number of generations elapsed in fragmentation conditions, the larger the negative magnitude of effect sizes on heterozygosity. Our results also suggest that fragmentation is shifting mating patterns towards increased selfing. We conclude that current conservation efforts in fragmented habitats should be focused on common or recently rare species and mainly outcrossing species and outline important issues that need to be addressed in future research on this area.

Factors affecting phenological patterns of bombacaceous trees in seasonal forests in Costa Rica and Mexico

We compared phenological patterns of tree species of the family Bombacaceae in three seasonal forests in Mexico and Costa Rica whose dry seasons vary in duration and intensity. The objectives were to (1) determine intraspecific variation in phenology between sites in different geographic locations with different precipitation regimes, (2) compare interspecific phenological patterns within sites during one year, and (3) document seasonal pollinator use of floral resources at one site in relation to the flowering phenology of these species. To determine the sequence of phenological events in trees of the family Bombacaceae across three study sites, phenology of marked individuals was recorded every 2 wk from September 2000 through August 2001 for six species. To estimate the importance of bombacaceous species in the diet of nectarivorous bats, pollen samples were collected from the bodies or feces of bats once every 2 wk during flowering. Our study suggests that phenological patterns of the Bombacaceae family in Neotropical dry forests are mainly constrained by phylogenetic membership and adaptive selective pressures associated with competition for pollinators. Abiotic factors related to precipitation and soil water content appear to be regulating leaf flushing and abscission, but the principal causes of flowering are related to ultimate factors associated with competition for pollinators. This study is the first that evaluates the phenological pattern of species and genera of the same family at different latitudes in a similar life zone.

Pollen load size, reproductive success, and progeny kinship of naturally pollinated flowers of the tropical dry forest tree Pachira quinata (Bombacaceae)

Several studies have demonstrated, using controlled pollinations, that the number and identity of pollen grains deposited onto a flowers stigma affect the reproductive success of plants. However, few studies have shown this relationship under conditions of natural pollination. Using the tropical dry forest tree Pachira quinata, we evaluated the relationship between the number of microgametophytes per pistil and the number of sires with respect to the production of fruits and seeds in a natural population of Pachira quinata. Our study demonstrates that fruit and seed production are directly related to the number of microgametophytes per pistil in natural populations of P. quinata. Only 6% of the marked flowers developed into mature fruits and 10% of the marked flowers initiated fruits but later aborted them. A mean of 23 pollen grains were required to produce a seed. Flowers with >400 pollen grains on the stigma always developed into mature fruits, whereas flowers that received <200 grains never matured fruits. Half of the pollen grains transferred to a flower stigma germinated and developed pollen tubes to the base of the style. The number of pollen grains on a stigma explained 34% of the variation in seed number per fruit, and the number of seeds produced per fruit is positively correlated with the size of the seeds. The population of P. quinata studied is predominantly outcrossing, and seeds within fruits are sired by one or a few donors. The total seed crop within trees was sired by three to five donors. Our study examined the implications of the variation in size of microgametophyte loads per pistil with respect to the breeding system and the paternity of progeny under natural conditions. The competitive ability of pollen and pollen tube attrition are important factors regulating fruit production in P. quinata.

The Weedy Rice Complex in Costa Rica. I. Morphological Study of Relationships Between Commercial Rice Varieties, Wild Oryza Relatives and Weedy Types

Weedy rice is a complex of Oryza morphotypes widely distributed in commercial rice fields, which interfere with rice cultivation, seed production, industrial processing and commercialization of this crop in several countries. The objective of this study was to characterize the weedy rice complex of Costa Rica by comparing it with the cultivated and wild rice species found in the country. A collection of weedy rice accessions, representative of the morphotypes found in the country, was established and characterized. Their morphometric relationships were established by comparing 27 morphological traits with commercial rice cultivars, landraces and wild Oryza species and by performing a multivariate analysis. Twenty-one weedy rice morphotypes were identified among 735 weedy accessions by using a three-digit code based on seed characters. Three principal components (PCs) explained 66.25% of the variation observed. The first PC accounted for 36.21% of the variation and separated CCDD genome type Oryza latifolia and O. grandiglumis from AA genome species O. sativa, O. glumaepatula, O. rufipogon and O. glaberrima. The second (18.9%) and third (11.14%) PCs separated the weedy morphotype groups from the AA genome species O.sativa, O. glaberrima and O. rufipogon. The weedy morphotypes were scattered between the indica commercial rice varieties, the cluster landraces–glaberrima and O. rufipogon. Additionally, a group of morphotypes showed intermediate characteristics between O. sativa and O. rufipogon, suggesting that hybridization could have taken place in the past between these species. None of the morphotypes collected in Costa Rica clustered with the allotetraploids CCDD species or O. glumaepatula.

Morphometric, isozymic and mitochondrial variability of Africanized honeybees in Costa Rica.

Wing morphometry, isozymic variation and mitochondrial RFLPs were studied in samples of feral Africanized bees collected at three different locations in Costa Rica, two of them in the lowlands (Guanacaste and Golfito regions) and the other in the Central Valley. These data revealed a predominantly African origin at all locations. However, significant heterogeneity was found between the two lowland sites and the Central Valley for some isozyme markers as well as for the RFLPs considered in this study. Both isozymic and mitochondrial DNA European markers were more frequent in the Central Valley. This could be the result of different opportunities for European gene introgression at the beginning of the migration of Africanized honeybees. It is shown that different gene frequencies of mitochondrial and nuclear European genetic markers may exist in Africanized bee populations, and that this may result from different selection pressures among colonies with African and European queens.

Maximum likelihood estimates of gene frequencies and racial admixture in Apis mellifera L. (africanized honeybees)

In order to obtain estimates of gene frequencies and other genetic parameters in Apis mellifera, samples are usually obtained from groups of haploid male (drones) and sterile female (workers) offspring, each group being descended from a multiply-mated single queen. A probabilistic model of segregation is presented which allows for maximum-likelihood estimates of genetic parameters in the reproductive individuals using the genotypic segregations of workers and drones from different colonies. Marginal probabilities of these segregations are calculated for each queen genotype by assuming a Dirichlet distribution for the fertilization frequencies of the different male alleles stored in the spermathecae. Estimates of gene frequencies and racial proportions in Africanized honey bees from different localities in Brazil and Central America reveal geographical heterogeneity in these parameters, possibly as a result of different levels of gene flow from European races.

Long-term effects of habitat fragmentation on mating patterns and gene flow of a tropical dry forest tree, Ceiba aesculifolia (Malvaceae: Bombacoideae)

UNLABELLED PREMISE OF THE STUDY Tropical forest loss and fragmentation isolate and reduce the size of remnant populations with negative consequences for mating patterns and genetic structure of plant species. In a 4-yr study, we determined the effect of fragmentation on mating patterns and pollen pool genetic structure of the tropical tree Ceiba aesculifolia in two habitat conditions: isolated trees in disturbed areas (≤3 trees/ha), and trees (≥6 trees/ha) in undisturbed mature forest. • METHODS Using six allozyme loci, we estimated the outcrossing rate (tm), the mean relatedness of progeny (rp) within and between fruits, the degree of genetic structure of pollen pools (Φft), and the effective number of pollen donors (Nep). • KEY RESULTS The outcrossing rates reflected a strict self-incompatible species. Relatedness of progeny within fruits was similar for all populations, revealing single sires within fruits. However, relatedness of progeny between fruits within trees was consistently greater for trees in fragmented conditions across 4 yr. We found high levels of genetic structure of pollen pools in all populations with more structure in isolated trees. The effective number of pollen donors was greater for trees in undisturbed forest than in disturbed conditions. • CONCLUSIONS Our study showed that the progeny produced by isolated trees in disturbed habitats are sired by a fraction of the diversity of pollen donors found in conserved forests. The foraging behavior of bats limits the exchange of pollen between trees, causing higher levels of progeny relatedness in isolated trees.

Isozyme diversity and analysis of the mating system of the wild rice Oryza latifolia Desv. in Costa Rica

Oryza latifolia is the most abundant and widespread wild relative of rice in the lowlands of Costa Rica. It also has the most diverse morphology. However, little is known about the genetic diversity and mating system of this allotetraploid species. Genetic analyses of nine populations from different life zones from the Pacific and Atlantic slopes were performed using six isozyme loci. Differences in genetic variability were observed among populations, of which Palo Verde and Santa Rosa were the most diverse. Most Atlantic populations clustered together, and a similar result was obtained with the Pacific populations. High levels of interpopulation diversity were observed while most populations were monomorphic for at least one genotype, suggesting little genetic flow within populations. Even in polymorphic populations no variation was observed within progeny groups, combined with Hardy-Weinberg disequilibria in most populations, commonly observed in autogamous species or in species with clonal reproduction. However, the high frequency of heterozygous-like patterns may suggest that the reproductive system of O. latifolia might be more complex. Future research may explain the genetic patterns as well as the reproductive biology of this species. The knowledge of the genetic diversity and mating system of O. latifolia could contribute to the implementation of genetic resource conservation strategies and gene flow analyses, as well as of breeding programs for rice improvement.

Curculionid beetles in aborted flower buds and immature fruits of Ceiba pentandra (Bombacaceae)

We determined the incidence of curculionid beetles of the genus Lonchophorus on aborted and not aborted flower buds and developing fruits of the tree Ceiba pentandra in southeastern Costa Rica. Beetle larvae were found in reproductive parts of all trees sampled trees. The frequency of beetle larvae was greater in aborted buds and immature fruits. A positive correlation between larvae development and flower bud development indicates that female oviposition occurred in an early flower developmental stage and time until bud abortion is variable. Weevil herbivory could be considered as one of the main factors that cause flower bud and fruit abortion in C. pentandra.ResumenDeterminamos la incidencia de curculiónidos del género Lonchophorus en botones florales y frutos inmaduros tanto abortados como no abortados en el árbol Ceiba pentandra en el SE de Costa Rica. Las larvas de curculiónidos fueron encontradas en todos los árboles muestreados. La frecuencia de larvas fue mayor en botones y frutos abortados. Una correlación positiva entre el desarrollo larval y el desarrollo del botón indica que la oviposición ocurre en una fase temprana de este y que el tiempo hasta el aborto es variable. La herbivoría por larvas de curculiónidos puede ser uno de los factores más importantes en el aborto de botones y frutos inmaduros en C. pentandra.

Phenological patterns and reproductive success of Ceiba pentandra (Bombacaceae) in tropical dry and wet forests of Costa Rica

Estudiamos los patrones fenologicos y el exito reproductivo de 103 arboles de Ceiba pentandra, localizados en el bosque tropical seco y en el bosque tropical humedo de la costa del Pacifico de Costa Rica. Para determinar los patrones fenologicos se anoto la caida de hojas y la produccion de flores y frutos cada dos semanas desde diciembre hasta marzo para todos los arboles marcados por un periodo de tres anos (2001, 2002 y 2003). Tambien se tomaron datos de la frecuencia de floracion y fructificacion para dos anos mas (2000 y 2004). Nuestros datos sugieren que los patrones fenologicos de arboles de C. pentandra se comportan como ciclos irregulares mas que como ciclos fijos de intervalos supraanuales y el tipo de bosque en el cual los arboles se ubican no esta teniendo un efecto decisivo ni en la probabilidad ni en la frecuencia de los ciclos reproductivos. La ausencia de patrones de autocorrelacion negativa en el exito reproductivo (no reproduccion, solamente floracion y fructificacion) en anos sucesivos nos sugiere que los ciclos de floracion y fructificacion para esta especie no corresponden a un modelo simple de limitacion de recursos. Nuestros resultados muestran que no existe una relacion entre el exito reproductivo y la periodicidad de los ciclos reproductivos para esta especie.