Recent studies have demonstrated the increasingly important role of lianas (woody vines) in forest regeneration, species diversity and ecosystem-level processes, particularly in the tropics. Mechanisms responsible for the maintenance of liana species diversity could yield new insights into the maintenance of overall species diversity. Lianas contribute to forest regeneration and competition, not only by competing directly with trees, but also by differentially affecting tree species and thus changing how trees compete among themselves. In addition, they contribute considerably to ecosystem-level processes, such as whole-forest transpiration and carbon sequestration. As the rate of tropical forest disturbance increases, they are likely to increase in relative abundance throughout the tropics and the importance of lianas to many aspects of forest dynamics will grow.

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The ecology of lianas and their role in forests

Considerable ffort has been spent documenting correlations between dioecy and various ecological and morphological traits for the purpose of testing hypotheses about conditions that favor dioecy. The data analyzed in these studies, with few exceptions, come from local floras, within which it was possible to contrast he subsets of dioecious and nondioecious taxa with regard to the traits in question. However, if there is a strong phylogenetic component o the presence or absence of dioecy, regional sampling may result in spurious associations. Here, we report results of a categorical multivariate analysis of the strengths ofvarious associations of dioecy with other traits over all flowering plants. Families were scored for presence of absence of monoecy or dioecy, systematic position, numbers of species and genera, growth forms, modes of pollination and dispersal, geographic distribution, and trophic status. Seven percent of angiosperm genera (959 of 13,500) contain at least some dioecious species, and ;6% of angiosperm species (14,620 of 240,000) are dioecious. The most consistent associations in the data set relate the presence of dioecy to monoecy, wind or water pollination, and climbing rowth. At both the family and the genus level, insect pollination is underrepresented among dioecious plants. At the family level, a positive correlation between dioecy and woody growth results primarily from the association between dioecy and climbing growth (whether woody or herbaceous) because neither the tree nor the shrub growth forms alone are consistently correlated with a family's tendency to include dioecious members. Dioecy appears to have evolved most frequently via monoecy, perhaps through divergent adjustments of floral sex ratios between individual plants. Monoecy itself is related to abiotic pollination and climbing rowth as revealed by multivariate analysis. Dioecy and monoecy are concentrated in the less advanced superorders of Thorne (1992) and subclasses of Cronquist (1988). The frequency of dioecy found in a local flora therefore flects the level of dioecy in its particular pool of families as much as, or more than, local selective factors. The positive associations of dioecy with abiotic pollination and monoecy are related to floral developmental nd morphological attributes, as is the negative association with bird and bat pollination; the positive association of dioecy with climbing growth is tentatively explained in terms of differential se ection for optimal resource allocation to sexual function. If rapid upward growth is at a premium in climbers and if fruit set at least temporarily inhibits growth or requires the production of thicker, more slowly growing stems to support heavy fruits, itmight be advantageous to postpone femaleness. Ifthe effect is strong, this may favor male plants.

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Dioecy and its correlates in the flowering plants

The maintenance of species diversity by treefall gaps is a long-standing paradigm in forest ecology. Gaps are presumed to provide an environment in which tree species of differing competitive abilities partition heterogeneous resources. The empirical evidence to support this paradigm, however, remains scarce, and some recent studies even suggest that gaps do not maintain the diversity of shade-tolerant species. Although there is evidence that gaps maintain the diversity of pioneer trees, most of this evidence comes from studies that did not make comparisons between gaps and intact forest sites (controls). Further, nearly all studies on the maintenance of diversity by gaps have ignored lianas, an important component of both old-world and neotropical forests. We tested the hypothesis that treefall gaps maintain shade-tolerant tree, pioneer tree, and liana species diversity in an old-growth forest on Barro Colorado Island (BCI), Panama. We compared the density and species richness of these guilds between paired gap and non-gap sites on both a per- area and a per-individual (per capita) basis. We found no difference in shade-tolerant tree density and species richness between the gap and non-gap sites. Both pioneer tree and liana density and species richness, however, were significantly higher in the gap than in the non- gap sites on both a per-area and a per-individual basis. These results suggest that gaps maintain liana species diversity and that this effect is not merely a consequence of increased density. Furthermore, our data confirm the long-held belief that gaps maintain pioneer tree species diversity. Because lianas and pioneer trees combined account for ;43% of the woody plant species on BCI, and in other forests, our results are likely to be broadly applicable and suggest that gaps play a strong role in the maintenance of woody species diversity.

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Treefall gaps and the maintenance of species diversity in a tropical forest

Tropical forests are experiencing large-scale structural changes, the most apparent of which may be the increase in liana (woody vine) abundance and biomass. Lianas permeate most lowland tropical forests, where they can have a huge effect on tree diversity, recruitment, growth and survival, which, in turn, can alter tree community composition, carbon storage and carbon, nutrient and water fluxes. Consequently, increasing liana abundance and biomass have potentially profound ramifications for tropical forest composition and functioning. Currently, eight studies support the pattern of increasing liana abundance and biomass in American tropical and subtropical forests, whereas two studies, both from Africa, do not. The putative mechanisms to explain increasing lianas include increasing evapotranspirative demand, increasing forest disturbance and turnover, changes in land use and fragmentation and elevated atmospheric CO2. Each of these mechanisms probably contributes to the observed patterns of increasing liana abundance and biomass, and the mechanisms are likely to be interrelated and synergistic. To determine whether liana increases are occurring throughout the tropics and to determine the mechanisms responsible for the observed patterns, a widespread network of large-scale, long-term monitoring plots combined with observational and manipulative studies that more directly investigate the putative mechanisms are essential.

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Increasing liana abundance and biomass in tropical forests: emerging patterns and putative mechanisms

In tropical forests, lianas (woody vines) are important structural parasites of trees. We assessed the effects of forest fragmentation, treefall disturbance, soils, and stand attributes on liana communities in central Amazonian rain forests. Over 27 500 liana stems (≥2 cm diameter at breast height [dbh]) were recorded in 27 1-ha plots in continuous forest and 42 plots in 10 forest fragments ranging from 1 to 100 ha in area. For each plot, an index of forest disturbance was determined from a 20-yr study of tree-community dynamics, and 19 soil-texture and chemistry parameters were derived from soil surface samples (top 20 cm). Liana abundance was 187–701 stems/ha, and liana aboveground dry biomass varied from 3.7 to 12.3 Mg/ha. Liana abundance increased significantly near forest edges and was significantly positively associated with forest disturbance and significantly negatively associated with tree biomass. Liana biomass was similarly associated with disturbance and tree biomass but also increased signifi...

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Alwyn H. Gentry

Papers

The distribution and evolution of climbing plants