Mortality of large trees and lianas following experimental drought in an Amazon forest

Severe drought episodes such as those associated with El Ni?o Southern Oscillation (ENSO) events influence large areas of tropical forest and may become more frequent in the future. One of the most important forest responses to severe drought is tree mortality, which alters forest structure, composition, carbon content, and flammability, and which varies widely. This study tests the hypothesis that tree mortality increases abruptly during drought episodes when plant-available soil water (PAW) declines below a critical minimum threshold. It also examines the effect of tree size, plant life form (palm, liana, tree) and potential canopy position (understory, midcanopy, overstory) on drought-induced plant mortality. A severe, four-year drought episode was simulated by excluding 60% of incoming throughfall during each wet season using plastic panels installed in the understory of a 1-ha forest treatment plot, while a 1-ha control plot received normal rainfall. After 3.2 years, the treatment resulted in a 38% increase in mortality rates across all stems >2 cm dbh. Mortality rates increased 4.5-fold among large trees (>30 cm dbh) and twofold among medium trees (10-30 cm dbh) in response to the treatment, whereas the smallest stems were less responsive. Recruitment rates did not compensate for the elevated mortality of larger-diameter stems in the treatment plot. Overall, lianas proved more susceptible to drought-induced mortality than trees or palms, and potential overstory tree species were more vulnerable than midcanopy and understory species. Large stems contributed to 90% of the pretreatment live aboveground biomass in both plots. Large-tree mortality resulting from the treatment generated 3.4 times more dead biomass than the control plot. The dramatic mortality response suggests significant, adverse impacts on the global carbon cycle if climatic changes follow current trends.     

Ethnobotany of Woody Species in Second-Growth, Old-Growth, and Selectively Logged Forests of Northeastern Costa Rica

Abstract: We assessed quantitatively the woody species used for timber, medicine, and other products in 10 tropical wet-forest stands with different land-use histories in the Atlantic lowlands of northeastern Costa Rica. Species were classified into 20 use categories based on regional ethnobotanical studies. Three size classes of woody vegetation were sampled in nested, contiguous plots along transects: trees (≥5 cm diameter at breast height [dbh]), saplings (>1 m high, <5 cm dbh), and seedlings (>20 cm high, <1 m high). Our study included five second-growth stands, three old-growth stands, and two selectively logged stands. Of the 459 woody species surveyed, 70% of the species and 86% of the total number of individuals had at least one use. Overall, species richness was highest for medicinal species (167 species). Absolute and relative abundance of medicinal and timber trees was significantly higher in second-growth stands than in old-growth and selectively logged stands. For 8 of the 15 use categories examined statistically, stem density showed no significant differences across forest types for any stem size class. Young, tropical, second-growth forests and selectively logged forests have high utilitarian as well as conservation value and will likely become important sources of forest products. The success of secondary forest regeneration, however, depends critically upon conservation of genetically diverse source populations in forest fragments and protected old-growth stands.     

Tree recruitment in an empty forest

To assess how the decimation of large vertebrates by hunting alters recruitment processes in a tropical forest, we compared the sapling cohorts of two structurally and compositionally similar forests in the Rio Manu floodplain in southeastern Peru. Large vertebrates were severely depleted at one site, Boca Manu (BM), whereas the other, Cocha Cashu Biological Station (CC), supported an intact fauna. At both sites we sampled small (> or =1 m tall, <1 cm dbh) and large (> or =1 cm and <10 cm dbh) saplings in the central portion of 4-ha plots within which all trees > or =10 cm dbh were mapped and identified. This design ensured that all conspecific adults within at least 50 m (BM) or 55 m (CC) of any sapling would have known locations. We used the Janzen-Connell model to make five predictions about the sapling cohorts at BM with respect to CC: (1) reduced overall sapling recruitment, (2) increased recruitment of species dispersed by abiotic means, (3) altered relative abundances of species, (4) prominence of large-seeded species among those showing depressed recruitment, and (5) little or no tendency for saplings to cluster closer to adults at BM. Our results affirmed each of these predictions. Interpreted at face value, the evidence suggests that few species are demographically stable at BM and that up to 28% are increasing and 72% decreasing. Loss of dispersal function allows species dispersed abiotically and by small birds and mammals to substitute for those dispersed by large birds and mammals. Although we regard these conclusions as preliminary, over the long run, the observed type of directional change in tree composition is likely to result in biodiversity loss and negative feedbacks on both the animal and plant communities. Our results suggest that the best, and perhaps only, way to prevent compositional change and probable loss of diversity in tropical tree communities is to prohibit hunting.     

Large ontogenetic declines in intra-crown leaf area index in two temperate deciduous tree species

The widespread occurrence of age-related changes in leaf morphology and allocation suggests that the leaf area index of individual trees (intra-crown LAI) may decline late in ontogeny. We used direct, within-canopy measurements to quantify the LAI of canopy trees with exposed crowns of two temperate deciduous species. Intra-crown LAI declined from approximately 7 to 4 in Acer saccharum, and from approximately 9.5 to 6.5 in Betula alleghaniensis, as tree size increased (from 15 to 72 cm diameter at breast height [dbh]). For A. saccharum, age (which varied from 30 to 160 years) was a significantly better predictor of LAI decline than dbh. We also modeled the effect of ontogenetic declines in LAI on understory light availability and found that light transmission increases significantly as canopy trees grow and mature. Our results thus suggest that gradual declines in LAI with tree age may play an important and overlooked role in contributing to the heterogeneity of sub-canopy light regimes in mature forests.     

Seasonal water stress tolerance and habitat associations within four neotropical tree genera

We investigated the relationship between habitat association and physiological performance in four congeneric species pairs exhibiting contrasting distributions between seasonally flooded and terra firme habitats in lowland tropical rain forests of French Guiana, including Virola and Iryanthera (Myristicaceae), Symphonia (Clusiaceae), and Eperua (Caesalpiniaceae). We analyzed 10-year data sets of mapped and measured saplings (stems >150 cm in height and <10 cm diameter at breast height [dbh]) and trees (stems > or =10 cm dbh) across 37.5 ha of permanent plots covering a 300-ha zone, within which seasonally flooded areas (where the water table never descends below 1 m) have been mapped. Additionally, we tested the response of growth, survival, and leaf functional traits of these species to drought and flood stress in a controlled experiment. We tested for habitat preference using a modification of the torus translation method. Strong contrasting associations of the species pairs of Iryanthera, Virola, and Symphonia were observed at the sapling stage, and these associations strengthened for the tree stage. Neither species of Eperua was significantly associated with flooded habitats at the sapling stage, but E. falcata was significantly and positively associated with flooded forests at the tree stage, and trees of E. grandiflora were found almost exclusively in nonflooded habitats. Differential performance provided limited explanatory support for the observed habitat associations, with only congeners of Iryanthera exhibiting divergent sapling survival and tree growth. Seedlings of species associated with flooded forest tended to have higher photosynthetic capacity than their congeners at field capacity. In addition, they tended to have the largest reductions in leaf gas exchange and growth rate in response to experimental drought stress and the least reductions in response to experimental inundation. The corroboration of habitat association with differences in functional traits and, to a lesser extent, measures of performance provides an explanation for the regional coexistence of these species pairs. We suggest that specialization to seasonally flooded habitats may explain patterns of adaptive radiation in many tropical tree genera and thereby provide a substantial contribution to regional tree diversity.     

Evaluation of some competition indices for the construction of Lebanon cedar (Cedrus libani A. Rich.) stand models

The purpose of this research is to test the precision of some published competition indices of Lebanon cedar (Cedrus libani A. Rich.) for the estimation of future periodic diameter increment of individual trees. Twenty- nine published competition indices were tested, using fifteen separate sets of data and their pooled values, collected from various stand age and site quality classes Lebanon cedar at Antalya. Temporary sample plots were taken in Elmali-Qamkuyusu (9 sample plots) and Finike-Pinarcik (6 sample plots) in 2001. Every plot was stem mapped (x and y coordinate system), diameter (dbh), total height, crown length, crown diameter and 10-year radial increment were recorded for trees greater than 4 cm in dbh. Then, in order to evaluate these competition indices for the prediction of the periodic diameter increment of the individual trees. Three linear models have been constructed for each competition index. It was found that the competition indices (Daniels et al., 1986; Biging and Dobbertin, 1995; Pukkala and Kolstr?m, 1987; Hegyi, 1974) with larger influence-zone areas produce better results.     

The patchiness of epifoliar fungi in tropical forests: host range, host abundance, and environment

Fungal symbioses affect the diversity, dynamics, and spatial patterns of trees in tropical forests. Their ecological importance is partly driven by their inherent patchiness. We used epifoliar fungi, a guild of common, benign, obligate, fungal symbionts of plants, as a model system to evaluate the relative importance of host phylogeny, host relative abundance, and microclimate on the three-dimensional distribution of plant-fungus symbioses. In parallel studies in rainforests in Panama and Australia, most epifoliar fungi were able to colonize several plant lineages but showed significant host preferences within the local plant community. More closely related plant species were not more likely to share fungal symbionts. Instead, fungal species were more likely to be shared by more abundant hosts, which supported a greater number and diversity of fungi. Environmental conditions strongly affected spatial distributions, with sites in the dark understory 2.5- to fourfold more likely to have epifoliar fungi than in the exposed forest canopy. In the understory, fungal incidence increased with canopy openness. Canopy trees supported only a subset of the fungal symbionts found in the understory, suggesting that adult trees are not reservoirs of these fungal symbionts for understory juveniles.     

Hotspots of Epiphytic Lichen Diversity in Two Young Managed Forests

Understanding within-stand variation in diversity of epiphytes will provide an improved basis for producing timber while conserving biological diversity. Two 80-ha, 50–year–old managed stands of conifers were surveyed to locate 0.4 ha putative "diversity" plots, the areas appearing most diverse in lichen epiphytes. These plots were generally located in areas made heterogeneous by canopy gaps, wolf trees (trees with large-diameter lower branches), and old-growth remnant trees. "Matrix" plots, in contrast, were chosen at random from the remaining, more homogenous forest. Diversity plots hosted from 25% to 40% more epiphytic lichen species than matrix plots in both stands. The strongest within-stand gradients in species composition were correlated with percentage of plot occupied by gaps and wolf trees. Percentage of the plot in gaps was correlated with species richness (r = 0.79). In the more structurally diverse stand, diversity and abundance of nitrogen-fixing "cyanolichens" were correlated with percentage of the plot occupied by gaps (0.5 < r < 0.9), and alectorioid lichens were correlated with percentage of the plot occupied by old-growth remnant trees (0.5 < r < 0.6). In the stand with more homogenous structure, percentage of the plot under gaps was correlated with regionally common species that were otherwise absent or sparse in the matrix. Protecting gaps, hardwoods, wolf trees, and old-growth remnant trees during thinning or other partial cutting is likely to promote the majority of epiphytic macrolichens in young conifer forests. Because these features are easily recognized on aerial photos and on the ground by land managers, it is practical to manage for forest structures that would promote lichen diversity.     

Rain forest fragmentation and the proliferation of successional trees

The effects of habitat fragmentation on diverse tropical tree communities are poorly understood. Over a 20-year period we monitored the density of 52 tree species in nine predominantly successional genera (Annona, Bellucia, Cecropia, Croton, Goupia, Jacaranda, Miconia, Pourouma, Vismia) in fragmented and continuous Amazonian forests. We also evaluated the relative importance of soil, topographic, forest dynamic, and landscape variables in explaining the abundance and species composition of successional trees. Data were collected within 66 permanent 1-ha plots within a large (approximately 1000 km2) experimental landscape, with forest fragments ranging from 1 to 100 ha in area. Prior to forest fragmentation, successional trees were uncommon, typically comprising 2-3% of all trees (> or =10 cm diameter at breast height [1.3 m above the ground surface]) in each plot. Following fragmentation, the density and basal area of successional trees increased rapidly. By 13-17 years after fragmentation, successional trees had tripled in abundance in fragment and edge plots and constituted more than a quarter of all trees in some plots. Fragment age had strong, positive effects on the density and basal area of successional trees, with no indication of a plateau in these variables, suggesting that successional species could become even more abundant in fragments over time. Nonetheless, the 52 species differed greatly in their responses to fragmentation and forest edges. Some disturbance-favoring pioneers (e.g., Cecropia sciadophylla, Vismia guianensis, V. amazonica, V. bemerguii, Miconia cf. crassinervia) increased by >1000% in density on edge plots, whereas over a third (19 of 52) of all species remained constant or declined in numbers. Species responses to fragmentation were effectively predicted by their median growth rate in nearby intact forest, suggesting that faster-growing species have a strong advantage in forest fragments. An ordination analysis revealed three main gradients in successional-species composition across our study area. Species gradients were most strongly influenced by the standlevel rate of tree mortality on each plot and by the number of nearby forest edges. Species-composition also varied significantly among different cattle ranches, which differed in their surrounding matrices and disturbance histories. These same variables were also the best predictors of total successional-tree abundance and species richness. Successional-tree assemblages in fragment interior plots (>150 m from edge), which are subjected to fragment area effects but not edge effects, did not differ significantly from those in intact forest, indicating that area effects per se had little influence on successional trees. Soils and topography also had little discernable effect on these species. Collectively, our results indicate that successional-tree species proliferate rapidly in fragmented Amazonian forests, largely as a result of chronically elevated tree mortality near forest edges and possibly an increased seed rain from successional plants growing in nearby degraded habitats. The proliferation of fast-growing successional trees and correlated decline of old-growth trees will have important effects on species composition, forest dynamics, carbon storage, and nutrient cycling in fragmented forests.     

Lemurs and the Regeneration of Dry Deciduous Forest in Madagascar

Abstract: We sought to assess the role of lemurs for seed dispersal in the dry deciduous forest of western Madagascar and the possible consequences of the demise of lemurs for forest regeneration. Forest regeneration was studied in eight plots in two large blocks of primary forest and in seven fragments of primary forest (1 plot per fragment). In 4 of the 15 study plots, the abundance of saplings was negatively and significantly correlated ( p < 0.05) with the abundance of mature individuals of the same tree species. In another 10 study plots there were negative correlations, although these were not significant on the community level. Second-order statistics were significant with p < 0.001 and indicated that seed dispersal away from the parent trees was important for successful establishment of saplings. Apart from possibly the bush pig ( Potamochoerus larvatus ), only one vertebrate species of the dry forest, the brown lemur ( Eulemur fulvus ), ingested seeds> 11 mm long and passed them through its digestive tract unharmed. These results for lemurs were based on direct observations and fecal analyses. To evaluate the role of E. fulvus , we compared regeneration in forest plots with and without E. fulvus . In forest fragments without E. fulvus , fewer lemur-dispersed tree species regenerated than would be expected based on the presence of mature tree species that are lemur-dispersed ( p < 0.05). No such effect was seen in primary forests with E. fulvus or for trees whose seeds can also be dispersed by other vertebrates. Thus, regeneration of the dry deciduous forest of western Madagascar with the complete set of primary forest tree species seems to depend upon the presence of E. fulvus .     

Architecture of 54 moist-forest tree species: traits, trade-offs, and functional groups

Tree architecture is an important determinant of the height extension, light capture, and mechanical stability of trees, and it allows species to exploit the vertical height gradient in the forest canopy and horizontal light gradients at the forest floor. Tropical tree species partition these gradients through variation in adult stature (Hmax) and light demand. In this study we compare 22 architectural traits for 54 Bolivian moist-forest tree species. We evaluate how architectural traits related to Hmax vary with tree size, and we present a conceptual scheme in which we combine the two axes into four different functional groups. Interspecific correlations between architecture and Hmax varied strongly from negative to positive, depending on the reference sizes used. Stem height was positively related to Hmax at larger reference diameters (14-80 cm). Species height vs. diameter curves often flattened toward their upper ends in association with reproductive maturity for species of all sizes. Thus, adult understory trees were typically shorter than similar-diameter juveniles of larger species. Crown area was negatively correlated with Hmax at small reference heights and positively correlated at larger reference heights (15-34 m). Wide crowns allow the small understory species to intercept light over a large area at the expense of a reduced height growth. Crown length was negatively correlated with Hmax at intermediate reference heights (4-14 m). A long crown enables small understory species to maximize light interception in a light-limited environment. Light-demanding species were characterized by orthotropic stems and branches, large leaves, and a monolayer leaf arrangement. They realized an efficient height growth through the formation of narrow and shallow crowns. Light demand turned out to be a much stronger predictor of tree architecture than Hmax, probably because of the relatively low, open, and semi-evergreen canopy at the research site. The existence of four functional groups (shade-tolerant, partial-shade-tolerant, and long- and short-lived pioneer) was confirmed by the principal component and discriminant analysis. Both light demand and Hmax capture the major variation in functional traits found among tropical rain forest tree species, and the two-way classification scheme provides a straightforward model to understand niche differentiation in tropical forests.     

Complementary Roles of Home Gardens and Exotic Tree Plantations as Alternative Habitats for Plants of the Ethiopian Montane Rainforest

Abstract:  Many tropical forests have been converted for agri- or silviculture or a combination of both (agroforestry). Conservation at a landscape scale requires an understanding of the distribution and abundance of native biodiversity in these converted natural ecosystems, of which the knowledge is especially poor for African agroecosystems. We compared species density and species composition of four plant groups (trees and shrubs, epiphytic vascular plants, mosses, and liverworts) among three arboreal land-use types in southwestern Ethiopia (montane rainforest fragments, shade-tree coffee home gardens, and exotic tree plantations). Species density was significantly higher in forests than in coffee home gardens for all plant groups and in exotic tree plantations for all groups except mosses. Home gardens had more vascular epiphytic species than plantations, whereas the reverse was true for mosses and liverworts. The species composition of the forest plots was sometimes more similar to home-garden plots than plantation plots and sometimes vice versa. Fifteen forest plots had, however, cumulatively more species than a random selection of 15 nonforest (coffee home garden and plantation) plots, even if the 2 plot types complemented each other in terms of habitats for forest plants. Tree plantations dominated by Eucalyptus had many small trees and shrubs in common with forests, whereas plantations with Cupressus were important substrates for forests mosses and liverworts. Our results illustrate the importance of undisturbed forests habitats for conservation of species at a landscape scale and that different human-made land-use types may complement each other in their capacity as additional habitats for forest species.     

Functional traits and the growth-mortality trade-off in tropical trees

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

Using forest inventory data to assess fisher resting habitat suitability in California.

The fisher (Martes pennanti) is a forest-dwelling carnivore whose current distribution and association with late-seral forest conditions make it vulnerable to stand-altering human activities or natural disturbances. Fishers select a variety of structures for daily resting bouts. These habitat elements, together with foraging and reproductive (denning) habitat, constitute the habitat requirements of fishers. We develop a model capable of predicting the suitability of fisher resting habitat using standard forest vegetation inventory data. The inventory data were derived from Forest Inventory and Analysis (FIA), a nationwide probability-based sample used to estimate forest characteristics. We developed the model by comparing vegetation and topographic data at 75 randomly selected fisher resting structures in the southern Sierra Nevada with 232 forest inventory plots. We collected vegetation data at fisher resting locations using the FIA vegetation sampling protocol and centering the 1-ha FIA plot on the resting structure. To distinguish used and available inventory plots, we used nonparametric logistic regression to evaluate a set of a priori biological models. The top model represented a dominant portion of the Akaike weights (0.87), explained 31.5% of the deviance, and included the following variables: average canopy closure, basal area of trees <51 cm diameter breast height (dbh), average hardwood dbh, maximum tree dbh, percentage slope, and the dbh of the largest conifer snag. Our use of routinely collected forest inventory data allows the assessment and monitoring of change in fisher resting habitat suitability over large regions with no additional sampling effort. Although models were constrained to include only variables available from the list of those measured using the FIA protocol, we did not find this to be a shortcoming. The model makes it possible to compare average resting habitat suitability values before and after forest management treatments, among administrative units, across regions and over time. Considering hundreds of plot estimates as a sample of habitat conditions over large spatial scales can bring a broad perspective, at high resolution, and efficiency to the assessment and monitoring of wildlife habitat.     

Effects of Selective Logging on Bat Communities in the Southeastern Amazon

Abstract:  Although extensive areas of tropical forest are selectively logged each year, the responses of bat communities to this form of disturbance have rarely been examined. Our objectives were to (1) compare bat abundance, species composition, and feeding guild structure between unlogged and low-intensity selectively logged (1–4 logged stems/ha) sampling grids in the southeastern Amazon and (2) examine correlations between logging-induced changes in bat communities and forest structure. We captured bats in understory and canopy mist nets set in five 1-ha study grids in both logged and unlogged forest. We captured 996 individuals, representing 5 families, 32 genera, and 49 species. Abundances of nectarivorous and frugivorous taxa (Glossophaginae, Lonchophyllinae, Stenodermatinae, and Carolliinae) were higher at logged sites, where canopy openness and understory foliage density were greatest. In contrast, insectivorous and omnivorous species (Emballonuridae, Mormoopidae, Phyllostominae, and Vespertilionidae) were more abundant in unlogged sites, where canopy foliage density and variability in the understory stratum were greatest. Multivariate analyses indicated that understory bat species composition differed strongly between logged and unlogged sites but provided little evidence of logging effects for the canopy fauna. Different responses among feeding guilds and taxonomic groups appeared to be related to foraging and echolocation strategies and to changes in canopy cover and understory foliage densities. Our results suggest that even low-intensity logging modifies habitat structure, leading to changes in bat species composition.     

Effects of Long-Term Ungulate Exclusion and Recent Alien Species Control on the Preservation and Restoration of a Hawaiian Tropical Dry Forest

Abstract: Although the destruction of tropical rain forests receives much attention, tropical dry forests are in general far more threatened and endangered. Eliminating grazing ungulates is often considered a key first step toward protecting these ecosystems, but few studies have investigated the long-term effects of this technique. We examined the effects of ungulate exclusion from a 2.3-ha native dry-forest preserve on the island of Hawaii by comparing its present flora to the flora of an adjacent area subjected to continuous grazing since the preserve was fenced over 40 years ago. Relative to this adjacent area, the fenced preserve contained a more diverse flora with substantially greater coverage of native overstory and understory species. Until recently, however, regeneration of native canopy trees within the preserve appears to have been thwarted by a dominant herbaceous cover of alien fountain grass (   Pennisetum setaceum ) and predation by alien rodent species. Our results indicate that although ungulate exclusion may be a necessary and critical first step, it is not sufficient to adequately preserve and maintain Hawaii's remaining tropical dry forest remnants. Our recent efforts to control the dominant alien species within the fenced preserve suggest that this practice may facilitate both the regeneration of native species and the colonization and potential invasion of new alien plants. Comparisons of seedlings of the dominant native canopy tree Diospyros sandwicensis growing in sites both dominated by and free of fountain grass suggested that fountain grass inhibits Diospyros seedling growth and photosynthesis but may increase survival if seedlings are protected from ungulates.     

Specialization and interaction strength in a tropical plant-frugivore network differ among forest strata

The degree of interdependence and potential for shared coevolutionary history of frugivorous animals and fleshy-fruited plants are contentious topics. Recently, network analyses revealed that mutualistic relationships between fleshy-fruited plants and frugivores are mostly built upon generalized associations. However, little is known about the determinants of network structure, especially from tropical forests where plants' dependence on animal seed dispersal is particularly high. Here, we present an in-depth analysis of specialization and interaction strength in a plant-frugivore network from a Kenyan rain forest. We recorded fruit removal from 33 plant species in different forest strata (canopy, midstory, understory) and habitats (primary and secondary forest) with a standardized sampling design (3447 interactions in 924 observation hours). We classified the 88 frugivore species into guilds according to dietary specialization (14 obligate, 28 partial, 46 opportunistic frugivores) and forest dependence (50 forest species, 38 visitors). Overall, complementary specialization was similar to that in other plant-frugivore networks. However, the plant-frugivore interactions in the canopy stratum were less specialized than in the mid- and understory, whereas primary and secondary forest did not differ. Plant specialization on frugivores decreased with plant height, and obligate and partial frugivores were less specialized than opportunistic frugivores. The overall impact of a frugivore increased with the number of visits and the specialization on specific plants. Moreover, interaction strength of frugivores differed among forest strata. Obligate frugivores foraged in the canopy where fruit resources were abundant, whereas partial and opportunistic frugivores were more common on mid- and understory plants, respectively. We conclude that the vertical stratification of the frugivore community into obligate and opportunistic feeding guilds structures this plant-frugivore network. The canopy stratum comprises stronger links and generalized associations, whereas the lower strata are composed of weaker links and more specialized interactions. Our results suggest that seed-dispersal relationships of plants in lower forest strata are more prone to disruption than those of canopy trees.     

A Century of Plant Species Loss from an Isolated Fragment of Lowland Tropical Rain Forest

An isolated 4-ha fragment of lowland tropical rain forest has been preserved in the Singapore Botanic Gardens since their founding in 1859. The Botanic Gardens'Jungle has recently had enumerated all woody stems 5 cm diameter at breast height (dbh) and larger and the complete vascular plant flora inventoried. This inventory can be compared with the historic record of the flora of the Gardens'Jungle obtained from the extensive collection of herbarium specimens dating back to the 1890s. Of the 448 historically recorded native species, 220 are still present. Ninety-four native species for which there were no historic records and 80 introduced species were also recorded in the recent inventory. The 50.9% loss of plant species richness over approximately the last century has not been distributed uniformly across plant life-form groups. Tree species have been less likely to go extinct than shrubs, climbers, or epiphytes. But half of the tree species present in 1994 were represented by only one or two individuals ≥ 5 cm dbh and larger. Individual longevity may be the major correlate with persistence of plant species in isolated forest fragments. Shade-tolerant understory shrubs (mostly Rubiaceae ) and rattans ( Palmae ) have been particularly prone to extinction. Some species have probably proliferated during the period of isolation. The tree Calophyllum ferrugineum currently constitutes one quarter of all woody stems. A group of climbers has become very common and covers large areas, probably inhibiting tree regeneration. We conclude that tiny fragments will act as refuges for tropical rain-forest plant species for decades, possibly even centuries after isolation but on their own they will not provide a permanent guarantee of the conservation of tropical biodiversity.     

Canopy Seed Banks as Time Capsules of Biodiversity in Pasture-Remnant Tree Crowns

Abstract: Tropical pastures present multiple barriers to tree regeneration and restoration. Relict trees serve as “regeneration foci” because they ameliorate the soil microclimate and serve as safe spots for dispersers. Here, we describe another mechanism by which remnant trees may facilitate pasture regeneration: the presence of seed banks in the canopy soil that accumulates from decomposing epiphytes within the crowns of mature remnant trees in tropical cloud forest pastures. We compared seed banks of canopy soils (histosols derived from fallen leaves, fruits, flower, and twigs of host trees and epiphytes, dead bryophytes, bark, detritus, dead animals, and microorganisms, and dust that accumulate on trunks and the upper surfaces of large branches) in pastures, canopy soils in primary forest trees, and soil on the forest floor in Monteverde, Costa Rica. There were 5211 epiphytic and terrestrial plant seeds in the three habitats. All habitats were dominated by seeds in a relatively small number of plant families, most of which were primarily woody, animal pollinated, and animal dispersed. The density of seeds on the forest floor was greater than seed density in either pasture‐canopy or forest‐canopy soils; the latter two did not differ. Eight species in 44 families and 61 genera from all of the habitats were tallied. There were 37 species in the pasture‐canopy soil, 33 in the forest‐canopy soil, and 57 on the forest floor. Eleven species were common to all habitats. The mean species richness in the pasture canopy was significantly higher than the forest canopy (F =83.38; p < 0.02). Nonmetric multidimensional scaling ordination revealed that the communities were distinct. Greenhouse experiments verified that many of these seeds were viable, with 29 taxa germinating (23 taxa in pruned mats [mimic of exposed conditions] and 16 taxa in control mats [intact conditions]) within 2 months of observation. Nearly half the species that germinated were characteristic of primary forests (primary forest samples, 19%; pasture samples, 29%). This supports the idea that canopy seed banks of pasture trees can function as time capsules by providing propagules that are removed in both space and time from the primary forest. Their presence may enhance the ability of pastures to regenerate more quickly, reinforcing the importance of trees in agricultural settings.