Mauricio Quesada

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.

A quantitative review of pollination syndromes: do floral traits predict effective pollinators?

The idea of pollination syndromes has been largely discussed but no formal quantitative evaluation has yet been conducted across angiosperms. We present the first systematic review of pollination syndromes that quantitatively tests whether the most effective pollinators for a species can be inferred from suites of floral traits for 417 plant species. Our results support the syndrome concept, indicating that convergent floral evolution is driven by adaptation to the most effective pollinator group. The predictability of pollination syndromes is greater in pollinator-dependent species and in plants from tropical regions. Many plant species also have secondary pollinators that generally correspond to the ancestral pollinators documented in evolutionary studies. We discuss the utility and limitations of pollination syndromes and the role of secondary pollinators to understand floral ecology and evolution.

Effects of Forest Fragmentation on Pollinator Activity and Consequences for Plant Reproductive Success and Mating Patterns in Bat-pollinated Bombacaceous Trees1

ABSTRACT Forest fragmentation and the resulting spatial isolation of tree species can modify the activity of pollinators and may have important implications for the reproductive success and mating systems of the plants they pollinate. The objectives of this study were to (1) evaluate the effect of forest fragmentation on pollinator activity in bat-pollinated bombacaceous trees and (2) determine the effects of forest fragmentation on reproductive success and mating systems of bombacaceous trees. We studied these parameters in three bombacaceous tree species in tropical seasonal forest of Chamela, Jalisco, Mexico, and Osa and Guanacaste, Costa Rica. For Ceiba aesculifolia, more visits were observed in fragments by both Glossophaga soricina and Leptonycteris curasoae. For Ceiba grandiflora, Musonycteris harrisoni visited flowers exclusively in forest and G. soricina visited more flowers in forest than in fragments; no difference was shown by L. curasoae. For Ceiba pentandra in Chamela, no differences were found in visitation by G. soricina between forest and fragments; L. curasoae visited significantly more flowers in forest. Ceiba pentandra received more visits by Phyllostomus discolor than G. soricina in Guanacaste, whereas no bat visitors were observed in Osa. Total mean flower production was greater in fragments than forest for C. aesculifolia, whereas no difference was observed for C. grandiflora. Fruit set was greater in forest than in fragments for C. grandiflora, whereas no difference was observed for C. aesculifolia. Outcrossing rates were high for C. aesculifolia and C. grandiflora in Chamela, and for C. pentandra in Guanacaste, independent of tree habitat, while C. pentandra in Osa showed a mixed-mating system. The effects of forest fragmentation on bat pollinators, plant reproductive success, and mating patterns varied depending upon the bombacaceous species. This variability was associated with the effects that forest fragmentation may have on differences in flowering patterns, bat foraging behavior, and plant self-incompatibility systems.

Changes in tree and liana communities along a successional gradient in a tropical dry forest in south-eastern Brazil

We investigated changes in species composition and structure of tree and liana communities along a successional gradient in a seasonally dry tropical forest. There was a progressive increase in tree richness and all tree structural traits from early to late stages, as well as marked changes in tree species composition and dominance. This pattern is probably related to pasture management practices such as ploughing, which remove tree roots and preclude regeneration by resprouting. On the other hand, liana density decreased from intermediate to late stages, showing a negative correlation with tree density. The higher liana abundance in intermediate stage is probably due to a balanced availability of support and light availability, since these variables may show opposite trends during forest growth. Predicted succession models may represent extremes in a continuum of possible successional pathways strongly influenced by land use history, climate, soil type, and by the outcomes of tree–liana interactions.

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.

INFLUENCE OF POLLINATION SPECIALIZATION AND BREEDING SYSTEM ON FLORAL INTEGRATION AND PHENOTYPIC VARIATION IN IPOMOEA

Natural selection should reduce phenotypic variation and increase integration of floral traits involved in placement of pollen grains on stigmas. In this study, we examine the role of pollinators and breeding system on the evolution of floral traits by comparing the patterns of floral phenotypic variances and covariances in 20 Ipomoea species that differ in their level of pollination specialization and pollinator dependence incorporating phylogenetic relatedness. Plants with specialized pollination (i.e., those pollinated by one functional group or by few morphospecies) displayed less phenotypic variation and greater floral integration than generalist plants. Self‐compatible species also displayed greater floral integration than self‐incompatible species. Floral traits involved in pollen placement and pick up showed less variation and greater integration than floral traits involved in pollinator attraction. Analytical models indicate that both breeding system and the number of morphospecies had significant effects on floral integration patterns although only differences in the former were significant after accounting for phylogeny. These results suggest that specialist/self‐compatible plants experience more consistent selection on floral traits than generalist/self‐incompatible plants. Furthermore, pollinators and breeding system promote integration of floral traits involved in pollen placement and pick up rather than integration of the whole flower.

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.

Effects of pollination by bats on the mating system of Ceiba pentandra (Bombacaceae) populations in two tropical life zones in Costa Rica

The identity and behavior of pollinators are among the main factors that determine the reproductive success and mating system of plants; however, few studies have directly evaluated the relationship between pollinators and the breeding system of the plants they pollinate. It is important to document this relationship because the global decline in pollinators may significantly affect the breeding systems of many animal-pollinated plants, particularly specialized systems. Ceiba pentandra is a tropical tree that has chiropterophilic flowers and a variable breeding system throughout its distribution, ranging from fully self-incompatible, to a mixed system with different degrees of selfing. To determine if regional differences in pollinators may result in regional differences in the outcrossing rate of this species, we used systematic observations of pollinator behavior in two tropical life zones and high-resolution genetic analysis of the breeding system of populations from these two regions using microsatellites. We found a predominantly self-incompatible system in regions with high pollinator visitation, while in environments with low pollinator visitation rates, C. pentandra changed to a mixed mating system with high levels of self-pollination.

Population dynamics, reproduction, and diet of the lesser long-nosed bat (Leptonycteris curasoae) in Jalisco, Mexico: implications for conservation

We estimate fluctuations in population size and sex ratio, documentbreeding behavior and reproduction, and determine the diet of a population ofthe lesser long-nosed bat, Leptonycteris curasoae, in anisland cave in Chamela Bay, Jalisco, Mexico, with monthly sampling during anannual cycle (October 1999–October 2000). Based on the area of thecaves ceiling and wall covered with L. curasoae inrelation to the potential roost area without them, in 1999 the abundanceincreased from 80% in October to 100% in November and December. In 2000 thepopulation decreased to 80% in January, 50% in February, 30% in March, 20% inApril, 10% in May, 5% in June and July, and less than 1% in August. Thepopulation rapidly increased to 60% in September and to 80% in October.Throughout the year there were significantly more males than females; however,there was significant heterogeneity over months. In September–Novemberthere were more females, but in December–August there were more malespresent. The majority of pregnant and lactating females were observed fromDecember to March and in July, while males were reproductive fromSeptember–January and in May–June. Breeding activity was observed inthe cave in November–December. Twenty-six species of plants were consumedduring the year, based on pollen identification from fecal samples. Bombacaceousspecies were the most important component of the diet from January to May andCactaceae were most important in June–September. Peak abundance and reproductive activitycoincided with peak flower resource availability, which occurred between Octoberand January and in June–July. The year-round presence and reproductiveactivity of L. curasoae at this site throughout the yeardemonstrate that many individuals are annual residents in this area and indicatethe importance of this roosting site. In order to develop a successfulconservation program for L. curasoae, in addition toprotecting migratory corridors and northern maternity roosts, it is equallyimportant to identify and protect areas that function as breeding colonies andyear-round sanctuaries for resident populations in the south.

Local and Landscape Factors Determining Occurrence of Phyllostomid Bats in Tropical Secondary Forests

Neotropical forests are being increasingly replaced by a mosaic of patches of different successional stages, agricultural fields and pasture lands. Consequently, the identification of factors shaping the performance of taxa in anthropogenic landscapes is gaining importance, especially for taxa playing critical roles in ecosystem functioning. As phyllostomid bats provide important ecological services through seed dispersal, pollination and control of animal populations, in this study we assessed the relationships between phyllostomid occurrence and the variation in local and landscape level habitat attributes caused by disturbance. We mist-netted phyllostomids in 12 sites representing 4 successional stages of a tropical dry forest (initial, early, intermediate and late). We also quantitatively characterized the habitat attributes at the local (vegetation structure complexity) and the landscape level (forest cover, area and diversity of patches). Two focal scales were considered for landscape characterization: 500 and 1000 m. During 142 sampling nights, we captured 606 individuals representing 15 species and 4 broad guilds. Variation in phyllostomid assemblages, ensembles and populations was associated with variation in local and landscape habitat attributes, and this association was scale-dependent. Specifically, we found a marked guild-specific response, where the abundance of nectarivores tended to be negatively associated with the mean area of dry forest patches, while the abundance of frugivores was positively associated with the percentage of riparian forest. These results are explained by the prevalence of chiropterophilic species in the dry forest and of chiropterochorous species in the riparian forest. Our results indicate that different vegetation classes, as well as a multi-spatial scale approach must be considered for evaluating bat response to variation in landscape attributes. Moreover, for the long-term conservation of phyllostomids in anthropogenic landscapes, we must realize that the management of the habitat at the landscape level is as important as the conservation of particular forest fragments.