The effect of habitat fragmentation on communities of mutualists: Amazonian ants and their host plants

The consequences of fragmentation for communities of mutualist partners are for the most part unknown; moreover, most studies addressing this issue have been conducted on plant-pollinator communities. We evaluated how the experimental fragmentation of lowland Amazonian rain forest influenced a community of ant-plant mutualists. We inventoried a total of 1057 myrmecophytes in four fragments and four continuous forest sites; the twelve plant species recorded were occupied by 33 ant morphospecies, of which 11 were obligate plant inhabitants. Neither plant species richness, ant species richness, nor total ant-plant density were significantly lower in forest fragments. However, eight of the plant species, including three of the four most common, had higher mean densities in continuous forest than fragments. Of these four species, only one (Cordia nodosa) had significantly different colonization rates between habitat types, with higher colonization rates of plants in fragments. This may be because the Azteca species it is associated with increases in abundance in forest isolates. Although our results suggest that communities of ant-plant mutualists are likely to persist in fragmented tropical landscapes 25 years after fragment isolation, most species are rare and populations sizes in fragments are extremely low. Environmental and demographic stochasticity could therefore limit long-term population viability. We suggest future studies focus on evaluating how fragmentation has altered herbivore pressure and the dispersal of ants and plants to fragments, since the interaction of these factors is likely to have the greatest impact on long-term patterns of population persistence.     

Habitat fragmentation reduces plant fitness by disturbing pollination and modifying response to herbivory

In fragmented landscapes plant species are often confined to remnants of formerly more widespread habitats, with many of their populations being small and isolated. This study experimentally examined the effects of population size and isolation on pollination, herbivory and reproductive success in the forest herb Phyteuma spicatum (Campanulaceae). In an experiment in which population size and isolation were manipulated using plants from the same origin, population size positively affected pollinator visitation, but did not alter the generally high levels of herbivory. As a result, seed production was higher in large populations. Conversely, plants originating from 14 natural populations of varying size and degree of isolation did not differ in reproductive success when grown in the same environment, suggesting similar attractiveness to pollinators and reproductive potential. The intensity of herbivory, however, was higher in progeny of small populations, at least in terms of the proportion of biomass removed. In both experiments, there were no effects of population isolation. The results suggest (1) that small population size decreases reproductive success via direct negative effects on plant-pollinator interactions, (2) that this pattern is not offset by herbivory, but (3) that herbivory enforces fragmentation effects on pollination by further reducing the number of flowering individuals and (4) that habitat fragmentation may influence plant fitness by affecting plant response to herbivory. The effects of habitat fragmentation on plant populations in present-day landscapes are thus complex, illustrating the need for more integrated studies in conservation biology that take into account both mutualistic and antagonistic plant-animal interactions.     

Effect of forest fragmentation on fruit and seed predation of the tropical dry forest tree Ceiba aesculifolia

Forests fragmentation reduces the density of natural plant populations forming patches of the remaining individuals. One of the biotic interactions that can be affected by forest fragmentation and is poorly studied is seed predation. We determined the effects of forest fragmentation on seed and fruit predation in Ceiba aesculifolia by comparing trees in continuous forest with trees in fragmented forest. We compared the following variables: (a) frequency of fruit predation by Collie’s squirrel (Sciurus colliaei) in each habitat; (b) frequency of the cotton-staining bug seed predator (Dysdercus, Orden Hemiptera) in each habitat; (c) the effect of seed predation on germination frequency and time; and (d) the effect of different life stages of Dysdercus on seed viability. In continuous habitat, 100% of the trees presented fruits with squirrel predation while only 34% of trees in fragmented habitats presented fruit predation. In continuous forest 27% of the trees contained fruits with the seed predator Dysdercus, while only 2% of the trees in fragmented forest presented Dysdercus. The initial weight of damaged seeds was greater than seeds that were not damaged indicating that seed predators select heavier seeds to feed upon. Frequency of seed germination was affected by different life stages; pre-adults decreased germination significantly more than nymphs and adults. Seed predation significantly increased the time it took for germination to occur. Our study shows that forest fragmentation significantly affects predation patterns of squirrels and cotton-staining bugs. Reduction of natural seed predators in forest fragments may have long-term consequences on forest structure and diversity.     

Fragmented but not isolated: Contribution of single trees, small patches and long-distance pollen flow to genetic connectivity for Gomortega keule, an endangered Chilean tree

Habitat fragmentation is a major threat to species survival worldwide due to genetic isolation, inbreeding depression, genetic drift and loss of adaptive potential. However the data on how gene-flow changes following habitat fragmentation is contradictory. If there is significant gene-flow between spatially isolated populations then limited conservation resources could be directed away from projects to ‘establish genetic connectivity’ and used to address other consequences of habitat fragmentation.This research focused on an endangered tree species Gomortega keule (Gomortegaceae) in a fragmented landscape in the Central Chile Biodiversity Hotspot and addressed three questions: (1) How far does pollen move between pollen donors and seed trees and what is the shape of the dispersal curve? (2) Do insect pollinators travel outside of forest patches? (3) Do small populations and single trees contribute to genetic connectivity across the landscape?Paternity analysis results show that G. keule’s insect pollinators travel outside of forest patches, over distances of 6 km, beyond the scale of population fragmentation or genetic structure. Pollen moved from small sites and single trees into large sites, as well as in the other direction, indicating these sites play a key role as functioning elements of the wider population and as stepping stones between sites. Fragmentation at the scale investigated has not led to genetic isolation, thus genetic connectivity per se is not a conservation priority. Other consequences of land-use change, specifically continuing habitat loss and population reduction, still threaten the survival of the species.     

Habitat fragmentation reduces genetic diversity and connectivity among toad populations in the Brazilian Atlantic Coastal Forest

Tropical rainforests are becoming increasingly fragmented and understanding the genetic consequences of fragmentation is crucial for conservation of their flora and fauna. We examined populations of the toad Rhinella ornata, a species endemic to Atlantic Coastal Forest in Brazil, and compared genetic diversity among small and medium forest fragments that were either isolated or connected to large forest areas by corridors. Genetic differentiation, as measured by F_ST, was not related to geographic distance among study sites and the size of the fragments did not significantly alter patterns of genetic connectivity. However, population genetic diversity was positively related to fragment size, thus haplotype diversity was lowest in the smallest fragments, likely due to decreases in population sizes. Spatial analyses of genetic discontinuities among groups of populations showed a higher proportion of barriers to gene flow among small and medium fragments than between populations in continuous forest. Our results underscore that even species with relatively high dispersal capacities may, over time, suffer the negative genetic effects of fragmentation, possibly leading to reduced fitness of population and cases of localized extinction.     

Habitat fragmentation alters the structure of dung beetle communities in the Atlantic Forest

Habitat loss and fragmentation have turned into the most important threats to biodiversity and ecosystem function worldwide. Here we investigate the effects of habitat fragmentation and drastic changes in tree communities on dung beetle richness and community structure. This study was carried out in a severely fragmented 670-km² forest landscape of the Atlantic Forest of north-eastern Brazil. Sampling was carried out in 19 forest fragments between September 2007 and March 2008 with the use of pitfall traps and flight interception traps. A total of 5893 individuals and 30 species of dung beetle were collected. Fragment area and isolation were the most significant explanatory variables for predictable and conspicuous changes in dung beetle species richness. Smaller and isolated fragments presented lower number of species, but fragments with lower tree species richness and lower proportion of shade-tolerant species were also considerably impoverished in terms of dung beetle species richness. The body mass of dung beetles were explained by fragment area and the percentage of emergent trees with smaller and less stratified fragments being dominated by small-bodied dung beetles. An ordination analysis segregated dung beetle communities between small fragments (<100 ha) and the control area. Seventy-seven percent of the species were recorded in the control area and 22% of all species were unique to this habitat. Our findings indicate that large fragments in the Atlantic Forest appear to consist in a sort of irreplaceable habitats for particular groups of dung beetle species, as well as for the integrity of their communities.     

Spatial and temporal effects on recruitment of an Afromontane forest tree in a threatened fragmented ecosystem

Tropical forest fragmentation affects animal and plant populations in different ways. For plants, early stages (seed to seedling) are more sensitive to habitat alteration than adults, and can shape their future spatial patterns. Therefore, studying how seed germination and seedling growth and survival vary at different spatiotemporal scales enhances our understanding about plant recruitment in fragmented ecosystems. In this study we examine if, and to what extent, recruitment at early life-stages of Xymalos monospora (Monomiaceae), a bird-dispersed Afrotropical tree, differs between and within forest fragments that vary in size, surrounding matrix and microhabitats. Three years of field experiments (2004-2006) in south-east Kenya, revealed that patterns of seed germination and seedling survival and growth were largely inconsistent, both in space and time. Recruitment was not consistently higher in larger or less disturbed fragments. At smaller spatial scales within forest fragments, recruitment was subject to high between-year variation too, with decreased germination in gaps only in the dry year of 2004. However, performance of seeds and seedlings was consistently better away from than under conspecific fruiting trees. Our results imply that fragmented tree populations of X. monospora may become age-structured, or ultimately go extinct, if recruitment fails in subsequent years. This may especially affect populations in small, disturbed forest fragments, where seed dispersal and buffering against stochastic processes are generally reduced. Exotic plantations bordering indigenous forest fragments may provide suitable conditions for native tree recruitment; hence, forest expansion through enrichment planting should be considered in future conservation plans.     

Growth of an understory herb is chronically reduced in Amazonian forest fragments

The biotic and abiotic changes associated with habitat fragmentation have been shown to have major consequences for plant recruitment and survivorship. However, few studies have quantified the growth of plants that persist in fragments. Over the course of a decade, we measured annual growth of 5200 individuals of the common understory herb Heliconiaacuminata (Heliconiaceae) in an experimentally fragmented Amazonian forest. We tested (A) whether annual growth rates were lower in fragments than in continuous forest, and (B) whether cumulative growth rates of plants that survived the entire period were lower in fragments. While mean annual growth rates were often lower in fragments, differences were not significant in any year. After 10 years, however, the cumulative effect was that plants in fragments were significantly smaller. This had a clear demographic consequence – plants in fragments produced fewer inflorescences than plants in continuous forest. Our results demonstrate that chronic reduced individual growth may be an important mechanism contributing to reduced population viability in fragmented forests, and that negative demographic consequences of fragmentation for plants can take years to manifest themselves.     

Genetic diversity and local population structure of fragmented populations of Trillium camschatcense (Trilliaceae)

Trillium camschatcense, a long-lived common woodland herb, has been experiencing intensive habitat fragmentation over the last century in eastern Hokkaido, Japan. We examined the genetic diversity and population genetic structure of 12 fragmented populations with different population sizes using allozyme electrophoresis. The percentage of polymorphic loci and mean number of alleles per locus were positively related to population size, probably due to the stochastic loss of rare alleles (frequency of q<0.1) in small populations. Populations with 350 flowering plants or fewer had lost almost all of their rare alleles. While the heterozygosity and inbreeding coefficient were not related to population size, some small populations showed relatively high inbreeding coefficients. In spite of the low genetic differentiation among overall populations (F_ST=0.130), local population structuring was recognized between the two geographically discontinuous population groups. Within groups, sufficient historical gene flow was inferred, whereas a low dispersal ability of this species and geographical separation could produce apparent differentiation between groups.