Effects of Local Disturbance of Tropical Forests on Frugivores and Seed Removal of a Small-Seeded Afrotropical Tree

Abstract:  Small-scale, local disturbance of tropical forests, for example from selective logging, is widespread, but its effects on biodiversity and ecosystem function have rarely been studied. In 3 East African tropical rainforests, we investigated the effect of different levels of local forest disturbance on the frugivore community and on tree visitation and fruit removal rates of the small-seeded tree Celtis durandii. We quantified birds and primates in little and heavily disturbed sites, distinguishing between forest specialists, forest generalists, and forest visitors. We quantified frugivorous tree visitors and seed removal rates of C. durandii trees in the same sites. Forest disturbance reduced the species richness and density of the frugivore community and of forest specialists. Frugivorous species and individuals visiting the study trees were reduced significantly, which led to a marginally significant decline in fruit removal by all frugivores and a significant reduction in removal by forest specialists. Reduction in fruit removal by forest specialists was not compensated for by increases in removal by forest generalists or visitors. Results did not differ among the 3 rainforests, which suggests they were consistent at a regional scale. So local forest disturbance led to a loss of frugivores and their seed removal services. This suggests that large-seeded tree species and trees with small fruits are losing seed dispersers. Thus, local forest disturbance appears to have a more general negative impact on frugivores and their seed dispersal services than anticipated previously.     

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.     

Effects of Artificial Roosts for Frugivorous Bats on Seed Dispersal in a Neotropical Forest Pasture Mosaic

Abstract:  In the Neotropics ongoing deforestation is producing open and heavily fragmented landscapes dominated by agriculture, mostly plantations and cattle pastures. After some time agriculture often becomes uneconomical and land is abandoned. Subsequent habitat regeneration may be slow because seed inputs are restricted by a lack of incentives—such as suitable roost sites—for seed dispersers to enter deforested areas. Increasing environmental awareness has fostered growing efforts to promote reforestation. Practical and cost-efficient methods for kick-starting forest regeneration are, however, lacking. We investigated whether artificial bat roosts for frugivorous bat species can attract these key seed dispersers to deforested areas, thereby increasing seed rain. We installed artificial bat roosts in a forest-pasture mosaic in the Costa Rican Atlantic lowlands and monitored bat colonization and seed dispersal. Colonization occurred within a few weeks of installation, and 10 species of bats occupied the artificial roosts. Five species of frugivorous or nectarivorous bats colonized artificial roosts permanently in both primary habitat and in deforested areas, in numbers similar to those found in natural roosts. Seed input around artificial roosts increased significantly. Sixty-nine different seed types, mostly of early-successional plant species, were transported by bats to artificial roosts in disturbed habitats. The installation of artificial bat roosts thus successfully attracted frugivorous bats and increased seed inputs into degraded sites. This method is likely to speed up early-vegetation succession, which in turn will attract additional seed dispersers, such as birds, and provide a microhabitat for seeds of mid- and late-successional plants. As well as supporting natural forest regeneration and bat conservation, this cost-efficient method can also increase environmental awareness among landowners.     

Potential effects of forest policies on terrestrial biodiversity in a multi-ownership province.

We used spatial simulation models to evaluate how current and two alternative policies might affect potential biodiversity over 100 years in the Coast Ranges Physiographic Province of Oregon. This 2.3-million-ha province is characterized by a diversity of public and private forest owners, and a wide range of forest policy and management objectives. We evaluated habitat availability for seven focal species representing different life histories. We also examined how policies affected old-growth stand structure, age distributions relative to the historical range of variability, and landscape patterns of forest types. Under the current policy scenario, the area of habitat for old-growth forest structure and associated species increased over time, the habitat for some early-successional associates remained stable, and the area of hardwood vegetation and diverse early-successional stages declined. The province is projected to move toward but not reach the historical range of variation of forest age classes that may have occurred under the wildfire regimes of the pre-Euroamerican settlement period. Ownership explained much of the pattern of biodiversity in the province, and under the current policy scenario, its effect increased over time as the landscape diverged into highly contrasting forest structures and ages. Patch type diversity declined slightly overall but declined strongly within ownerships. Most of the modeled change in biodiversity over time resulted from policies on public forest lands that were intended to increase the area of late-successional forests and species. One of the alternative policies, increased retention of wildlife trees on private lands, reduced the contrast between ownerships and increased habitat availability over time for both early- and late-successional species. Analysis of another alternative, stopping thinning of plantations on federal lands, indicated that current thinning regimes improve habitat for the Olive-sided Flycatcher, but the no-thinning alternative had no effect on the habitat scores for the late-successional species in the 100-year simulation. A comparison of indicators of biological diversity suggests that using focal species and forest structural measures can provide complementary information on biodiversity. The multi-ownership perspective provided a more complete synthesis of province-wide biodiversity patterns than assessments based on single ownerships.     

Fragmentation drives tropical forest fragments to early successional states: A modelling study for Brazilian Atlantic forests

Land use leads to massive habitat destruction and fragmentation in tropical forests. Despite its global dimensions the effects of fragmentation on ecosystem dynamics are not well understood due to the complexity of the problem. We present a simulation analysis performed by the individual-based model FORMIND. The model was applied to the Brazilian Atlantic Forest, one of the world's biodiversity hot spots, at the Plateau of São Paulo. This study investigates the long-term effects of fragmentation processes on structure and dynamics of different sized remnant tropical forest fragments (1-100 ha) at community and plant functional type (PFT) level. We disentangle the interplay of single effects of different key fragmentation processes (edge mortality, increased mortality of large trees, local seed loss and external seed rain) using simulation experiments in a full factorial design.Our analysis reveals that particularly small forest fragments below 25 ha suffer substantial structural changes, biomass and biodiversity loss in the long term. At community level biomass is reduced up to 60%. Two thirds of the mid- and late-successional species groups, especially shade-tolerant (late successional climax) species groups are prone of extinction in small fragments. The shade-tolerant species groups were most strongly affected; its tree number was reduced more than 60% mainly by increased edge mortality. This process proved to be the most powerful of those investigated, explaining alone more than 80% of the changes observed for this group. External seed rain was able to compensate approximately 30% of the observed fragmentation effects for shade-tolerant species.Our results suggest that tropical forest fragments will suffer strong structural changes in the long term, leading to tree species impoverishment. They may reach a new equilibrium with a substantially reduced subset of the initial species pool, and are driven towards an earlier successional state. The natural regeneration potential of a landscape scattered with forest fragments appears to be limited, as external seed rain is not able to fully compensate for the observed fragmentation-induced changes. Our findings suggest basic recommendations for the management of fragmented tropical forest landscapes.     

Influence of environment, disturbance, and ownership on forest vegetation of coastal Oregon.

Information about how vegetation composition and structure vary quantitatively and spatially with physical environment, disturbance history, and land ownership is fundamental to regional conservation planning. However, current knowledge about patterns of vegetation variability across large regions that is spatially explicit (i.e., mapped) tends to be general and qualitative. We used spatial predictions from gradient models to examine the influence of environment, disturbance, and ownership on patterns of forest vegetation biodiversity across a large forested region, the 3-million-ha Oregon Coast Range (USA). Gradients in tree species composition were strongly associated with environment, especially climate, and insensitive to disturbance, probably because many dominant tree species are long-lived and persist throughout forest succession. In contrast, forest structure was strongly correlated with disturbance and only weakly with environmental gradients. Although forest structure differed among ownerships, differences were blurred by the presence of legacy trees that originated prior to current forest management regimes. Our multi-ownership perspective revealed biodiversity concerns and benefits not readily visible in single-ownership analyses, and all ownerships contributed to regional biodiversity values. Federal lands provided most of the late-successional and old-growth forest. State lands contained a range of forest ages and structures, including diverse young forest, abundant legacy dead wood, and much of the high-elevation true fir forest. Nonindustrial private lands provided diverse young forest and the greatest abundance of hardwood trees, including almost all of the foothill oak woodlands. Forest industry lands encompassed much early-successional forest, most of the mixed hardwood-conifer forest, and large amounts of legacy down wood. The detailed tree- and species-level data in the maps revealed regional trends that would be masked in traditional coarse-filter assessment. Although abundant, most early-successional forests originated after timber harvest and lacked legacy live and dead trees important as habitat and for other ecological functions. Many large-conifer forests that might be classified as old growth using a generalized forest cover map lacked structural features of old growth such as multilayered canopies or dead wood. Our findings suggest that regional conservation planning include all ownerships and land allocations, as well as fine-scale elements of vegetation composition and structure.     

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 .     

Successional changes in plant resistance and tolerance to herbivory

Despite considerable research on plant defenses, we know very little about how temporal changes in the environment may influence resistance and tolerance levels, or the costs and benefits of these defense strategies for long-lived plant species. We hypothesized that, in successional habitats, predictable environmental changes should favor strong plasticity in defense phenotypes and that the costs, benefits, and levels of tolerance and resistance will change with environmental context. Using a widely distributed, old-field perennial, late goldenrod (Solidago altissima), we conducted a field experiment to test these predictions. We planted goldenrod genets exhibiting varying levels of resistance and tolerance into three early-successional and three late-successional fields (approximately three and 15 years in age, respectively) and experimentally measured resistance and tolerance levels and their associated costs and selection coefficients. We found a significant effect of successional stage but no effect of genotype or stage-genotype interaction on defense levels. Genets planted in early-successional fields appeared to be more resistant and less tolerant to herbivory than those same genets planted in late-successional fields. There were significant trade-offs between resistance and tolerance in early-successional fields but not in late-successional fields. Each late-successional field exhibited a significant cost or selection gradient for resistance, but there was no general pattern of resistance costs or selection gradients specific to a successional stage class. In contrast, there was evidence of stage-specific costs of tolerance; late-successional fields exhibited significant costs of tolerance whereas early-successional fields did not. There was no evidence of direct selection for or against tolerance in either stage. Our results suggest that defense phenotypes might change in qualitative ways during succession. High resistance in early stages may be attributed to associational effects of the early-successional community, reducing the probability of damage, and despite a cost of tolerance in late stages, tolerance may be beneficial in mitigating the effects of both herbivory and environmental stresses (i.e., low light availability) that limit fitness in these fields. This study provides experimental evidence that succession can strongly influence defense phenotypes and promote temporal variability in relative resistance and tolerance levels.     

Survival and growth of under-planted trees: a meta-analysis across four biomes.

The transformation of natural forest regeneration processes by human activities has created the need to develop and implement new models of forest management. Alternative silvicultural systems such as variable retention harvest, partial and patch cuts, and older forest management practices such as under-planting, are used in many forests around the world, particularly in North American oak stands, the boreal and coastal temperate rain forests of Canada and the United States, and in many degraded tropical regions of Asia and the Americas. Specific objectives are pursued in each of these biomes, but some are common to most regions, such as preservation of cover and structure and their associated benefits for natural or artificial regeneration due to moderation of the microclimate, development of optimal light and competition conditions, and reduced predation by herbivores. Shelterwoods are often presented as an alternative to clear-cutting to improve the survival of planted trees. A meta-analysis of published results with randomization tests was performed to test the relationship between overstory density and planted seedling growth and survival. Multiple comparisons were also used to reveal optimal levels of overstory density, if they exist. A majority of studies show that survival and growth improve as stand density decreases to an intermediate level, below which they either drop or stabilize. This level seems optimal in most conditions, as it is also more apt to fulfill other objectives imposed on today's forest activities, such as the conservation of forest processes and structures, and the reconstruction of degraded stands through the accelerated return of mid- to late-successional species.     

Bat- and Bird-Generated Seed Rains at Isolated Trees in Pastures in a Tropical Rainforest

Abstract: Bats are abundant and effective seed dispersers inside the forest, but what happens when a forest is fragmented and transformed into pasture? The landscape at Los Tuxtlas, Mexico, originally rainforest, is greatly fragmented and covered with pastures. We analyzed the seed rains produced by frugivorous bats and birds under isolated trees in pastures in the fragmented landscape and the contribution of this process to vegetational recovery. We surveyed bats and obtained fecal samples under isolated trees in pastures. We also collected seed rain below the canopy of 10 isolated Ficus trees, separating nocturnally dispersed seeds from diurnally dispersed seeds. We caught 652 bats of 20 species; 83% of captures were frugivores. The most abundant species were Sturnira lilium (48%), Artibeus jamaicensis (18%), Carollia perspicillata (12%), and Dermanura tolteca (11%). Fecal samples contained seeds of 19 species in several families: Piperaceae (50%), Moraceae (25%), Solanaceae (12%), Cecropiaceae (10%), and others (3%). Sturnira lilium was the most important disperser bat in pastures. Seed rain was dominated by zoochorous species (89%). We found seed diversity between day and night seed captures to be comparable, but we found a significant interaction of disperser type (  bird or bat) with season. Seven plant species accounted for 79% of the seed rain: Piper auritum (23%), Ficus (  hemiepiphytic-strangler tree) spp. (17%), Cecropia obtusifolia (10%), P. amalago (10%), Ficus (  free-standing tree) spp. (8%), P. yzabalanum (6%), and Solanum rudepanum (5%). Bats and birds are important seed dispersers in pastures because they disperse seeds of pioneer and primary species (trees, shrubs, herbs, and epiphytes), connect forest fragments, and maintain plant diversity. Consequently, they might contribute to the recovery of woody vegetation in disturbed areas in tropical humid forests.     

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.     

More asymmetric tree competition brings about more evapotranspiration and less runoff from the forest ecosystems: A simulation study

The present paper reports how stand size-structure dynamics due to competition between different-sized trees affect long-term forested water balance in Japanese cool-temperate planted stands (evergreen coniferous Cryptomeria japonica and deciduous coniferous Larix kaempferi stands) using a fully coupled multi-layered meteorological surface physics—terrestrial ecosystems model. The simulation captured the well-known annual variation in leaf area index (LAI) accurately with stand age in monocultured and even-aged stands; the occurrence of maximum LAI during the early growth stage and then a gradual decline followed by a steady state after the maximum LAI. The simulations also detected a high dependency of annual evapotranspiration (AETr) on LAI with stand age that is well known by prior observational researches. In the C. japonica (shade-tolerant late-successional species) stand, the relationship between annual net primary productivity of an individual tree (NPP_ind) and individual tree mass (w) changed from linear to a convex curve during self-thinning, indicating that the degree of asymmetric tree competition intensified with forest stand development. The higher degree of competitive asymmetry characterized by the convex-shaped NPPind-w relationship produced greater size inequality, i.e., the formation of trees stratified by height. Under such conditions, AETr and annual transpiration (ATr) were mainly regulated by larger trees. On the other hand, the NPPind-w relationships in the L. kaempferi (shade-intolerant early-successional species) stand were linear throughout the simulated period, indicating the lower degree of competitive asymmetry. Under such conditions, the growth of intermediate-sized trees was enhanced and these trees became a dominant source of AETr (and also ATr) during self-thinning. Furthermore, the sensitivity analysis of the effects of ecophysiological parameters such as foliage profile (i.e., vertical distribution of leaf area density) of an individual tree (distribution pattern is described by the parameter η), the maximum carboxylation velocity (V_cmax0) and biomass allocation pattern of individual plant growth (μ₁) on AETr, ATr and annual runoff (AR_off) showed that the temporal trends of AETr, ATr, AR_off and NPPind-w relationships were completely the same as those in the control simulations. However, the NPPind-w relationship during self-thinning indicated higher degrees of competitive asymmetry when η or V_cmax0 were greater than those in the control simulation and generated greater AETr and ATr and thus smaller AR_off. We found that more asymmetric tree competition brings about greater size inequality between different-sized trees and thus more evapotranspiration and less runoff in a forest stand. Overall, our simulation approach revealed that not only LAI dynamics but also plant competition, and thus size-structure dynamics, in a forest ecosystem are essential to long-term future projections of forested water balance.     

Effects of Pioneer Tree Species Hyperabundance on Forest Fragments in Northeastern Brazil

Abstract: Despite many studies on fragmentation of tropical forests, the extent to which plant and animal communities are altered in small, isolated forest fragments remains obscure if not controversial. We examined the hypothesis that fragmentation alters the relative abundance of tree species with different vegetative and reproductive traits. In a fragmented landscape (670 km²) of the Atlantic Forest of northeastern Brazil, we categorized 4056 trees of 182 species by leafing pattern, reproductive phenology, and morphology of seeds and fruit. We calculated relative abundance of traits in 50 1‐ha plots in three types of forest configurations: forest edges, small forest fragments (3.4–83.6 ha), and interior of the largest forest fragment (3500 ha, old growth). Although evergreen species were the most abundant across all configurations, forest edges and small fragments had more deciduous and semideciduous species than interior forest. Edges lacked supra‐annual flowering and fruiting species and had more species and stems with drupes and small seeds than small forest fragments and forest interior areas. In an ordination of species similarity and life‐history traits, the three types of configurations formed clearly segregated clusters. Furthermore, the differences in the taxonomic and functional (i.e., trait‐based) composition of tree assemblages we documented were driven primarily by the higher abundance of pioneer species in the forest edge and small forest fragments. Our work provides strong evidence that long‐term transitions in phenology and seed and fruit morphology of tree functional groups are occurring in fragmented tropical forests. Our results also suggest that edge‐induced shifts in tree assemblages of tropical forests can be larger than previously documented.     

Tropical forest restoration: tree islands as recruitment foci in degraded lands of Honduras.

Tropical forest recovery in pastures is slowed by a number of biotic and abiotic factors, including a lack of adequate seed dispersal and harsh microclimatic extremes. Accordingly, methods to accelerate forest recovery must address multiple impediments. Here, we evaluated the ability of "tree islands" to serve as "recruitment foci" in a two-year study at three sites in northern Honduras. Islands of three sizes (64, 16, and 4 m2) and at two distances to secondary forest (20 and 50 m) were created by planting 2 m tall vegetative stakes of two native species: Gliricidia sepium (Fabaceae) and Bursera simaruba (Burseraceae), each in monoculture. Open-pasture "islands" of equal sizes served as controls. Tree islands reduced temperature and light (PAR) extremes as compared to open pasture, creating a microenvironment more favorable to seedling establishment. Seed-dispersing birds (quantified at one site only) showed an overwhelming preference for islands; 160 visits were recorded to islands compared with one visit to open pasture. Additionally, frugivores visited large islands more often, and for longer time periods, than small islands, thereby increasing the likelihood of a dispersal event there. In total, 144 140 seeds belonging to 186 species were collected in islands; more than 80% were grasses. Tree islands increased zoochorous tree seed rain; seed density and species richness were greater in tree islands than in open pasture, and large islands had greater seed density than smaller islands (Gliricidia only), suggesting that they are more effective for restoration. Distance to forest did not affect seed rain. A total of 543 seedlings and 41 species established in islands; > 85% were zoochorous. Seedling density did not differ among treatments (mean 0.2 seedlings/m2 for islands vs. 0.1 seedlings/m2 for pasture), although an increasing trend in tree islands over the course of two years suggests that seedling recruitment is accelerated there. Lastly, similar seedling densities were censused in the 1 m perimeter surrounding islands, suggesting that islands can expand outward into pasture. Planting vegetative stakes to create tree islands in pastures accelerates forest recovery by overcoming a number of impediments, and presents a simple, broadly applicable alternative for facilitating forest regeneration in abandoned pastures.     

Incorporating differences between genetic diversity of trees and herbaceous plants in conservation strategies

Reviews that summarize the genetic diversity of plant species in relation to their life history and ecological traits show that forest trees have more genetic diversity at population and species levels than annuals or herbaceous perennials. In addition, among-population genetic differentiation is significantly lower in trees than in most herbaceous perennials and annuals. Possible reasons for these differences between trees and herbaceous perennials and annuals have not been discussed critically. Several traits, such as high rates of outcrossing, long-distance pollen and seed dispersal, large effective population sizes (N_e), arborescent stature, low population density, longevity, overlapping generations, and occurrence in late successional communities, may make trees less sensitive to genetic bottlenecks and more resistant to habitat fragmentation or climate change. We recommend that guidelines for genetic conservation strategies be designed differently for tree species versus other types of plant species. Because most tree species fit an LH scenario (low [L] genetic differentiation and high [H] genetic diversity), tree seeds could be sourced from a few populations distributed across the species’ range. For the in situ conservation of trees, translocation is a viable option to increase N_e. In contrast, rare herbaceous understory species are frequently HL (high differentiation and low diversity) species. Under the HL scenario, seeds should be taken from many populations with high genetic diversity. In situ conservation efforts for herbaceous plants should focus on protecting habitats because the typically small populations of these species are vulnerable to the loss of genetic diversity. The robust allozyme genetic diversity databases could be used to develop conservation strategies for species lacking genetic information. As a case study of reforestation with several tree species in denuded areas on the Korean Peninsula, we recommend the selection of local genotypes as suitable sources to prevent adverse effects and to insure the successful restoration in the long term.     

Nonredundancy in the dispersal network of a generalist tropical forest tree

Plant species with generalized dispersal mutualisms are considered to be robust to local frugivore extinctions because of redundancy between dispersal agents. However, real redundancy can only occur if frugivores have similar foraging and ranging patterns and if fruit is a limiting resource. We evaluated the quantitative and qualitative contributions of seed dispersers for an endochorus mast-fruiting species, Prunus javanica (Rosaceae) in Khao Yai National Park, Thailand, to evaluate the potential redundancy of dispersers. Data were collected from tree watches, seed/fruit traps, and seed transects under and away from fruiting trees, feeding and seed deposition by gibbons (Hylobates lar), and evaluations of seed and first-year seedling survival. We identified three clusters of dispersers within the network. Most (>80%) frugivore species observed were small birds and squirrels that were not functional dispersers, dropping most seeds under or very near the tree crown, where seedling survival was ultimately nil. Monkeys (Macaca leonina) were low-quality, short-range dispersers, but they dispersed large numbers of seeds and were responsible for 67% of surviving first-year seedlings. Gibbons and Oriental Pied Hornbills (Anthracoceros albirostris) handled few fruits, but they provided the highest quality service by carrying most seeds away from the canopy to medium and long distances, respectively. Although there was overlap in the deposition patterns of the functional dispersers, they displayed complementary, rather than redundant, roles in seed dispersal. Satiation of all functional dispersers further limited their capacity to "replace" one another. Redundancy must be evaluated at the community level because each type of disperser may shift to different species in the non-masting years of P. javanica. Our results underscore the need for research on broader spatial and temporal scales, which combines studies of dispersal and plant recruitment, to better understand mechanisms that maintain network stability.     

Functional nonredundancy of elephants in a disturbed tropical forest

Conservation efforts are often motivated by the threat of global extinction. Yet if conservationists had more information suggesting that extirpation of individual species could lead to undesirable ecological effects, they might more frequently attempt to protect or restore such species across their ranges even if they were not globally endangered. Scientists have seldom measured or quantitatively predicted the functional consequences of species loss, even for large, extinction‐prone species that theory suggests should be functionally unique. We measured the contribution of Asian elephants (Elephas maximus) to the dispersal of 3 large‐fruited species in a disturbed tropical moist forest and predicted the extent to which alternative dispersers could compensate for elephants in their absence. We created an empirical probability model with data on frugivory and seed dispersal from Buxa Tiger Reserve, India. These data were used to estimate the proportion of seeds consumed by elephants and other frugivores that survive handling and density‐dependent processes (Janzen‐Connell effects and conspecific intradung competition) and germinate. Without compensation, the number of seeds dispersed and surviving density‐dependent effects decreased 26% (Artocarpus chaplasha), 42% (Careya arborea), and 72% (Dillenia indica) when elephants were absent from the ecosystem. Compensatory fruit removal by other animals substantially ameliorated these losses. For instance, reductions in successful dispersal of D. indica were as low as 23% when gaur (Bos gaurus) persisted, but median dispersal distance still declined from 30% (C. arborea) to 90% (A. chaplasha) without elephants. Our results support the theory that the largest animal species in an ecosystem have nonredundant ecological functionality and that their extirpation is likely to lead to the deterioration of ecosystem processes such as seed dispersal. This effect is likely accentuated by the overall defaunation of many tropical systems.     

The Normative Dimension and Legal Meaning of Endangered and Recovery in the U.S. Endangered Species Act

Abstract:  The ethical, legal, and social significance of the U.S. Endangered Species Act of 1973 (ESA) is widely appreciated. Much of the significance of the act arises from the legal definitions that the act provides for the terms threatened species and endangered species. The meanings of these terms are important because they give legal meaning to the concept of a recovered species. Unfortunately, the meanings of these terms are often misapprehended and rarely subjected to formal analysis. We analyzed the legal meaning of recovered species and illustrate key points with details from "recovery" efforts for the gray wolf ( Canis lupus ). We focused on interpreting the phrase "significant portion of its range," which is part of the legal definition of endangered species. We argue that recovery and endangerment entail a fundamentally normative dimension (i.e., specifying conditions of endangerment) and a fundamentally scientific dimension (i.e., determining whether a species meets the conditions of endangerment). Specifying conditions for endangerment is largely normative because it judges risks of extinction to be either acceptable or unacceptable. Like many other laws that specify what is unacceptable, the ESA largely specifies the conditions that constitute unacceptable extinction risk. The ESA specifies unacceptable risks of extinction by defining endangered species in terms of the portion of a species' range over which a species is "in danger of extinction." Our analysis indicated that (1) legal recovery entails much more than the scientific notion of population viability, (2) most efforts to recover endangered species are grossly inadequate, and (3) many unlisted species meet the legal definition of an endangered or threatened species.     

Effects of howler monkey reintroduction on ecological interactions and processes

Rewilding has been an increasingly popular tool to restore plant–animal interactions and ecological processes impaired by defaunation. However, the reestablishment of such processes has seldom been assessed. We investigated the restoration of ecological interactions following the reintroduction of the brown howler monkey (Alouatta guariba) to a defaunated Atlantic forest site. We expected the reintroduction to restore plant–animal interactions and interactions between howlers and dung beetles, which promote secondary seed dispersal. We estimated the number of interactions expected to be restored by the reintroduction to provide the baseline interaction richness that could be restored. We followed the reintroduced howler monkeys twice a week for 24 months (337 hours total) to assess their diet. We used howler monkey dung in secondary seed dispersal experiments with 2484 seed mimics to estimate the removal rates by dung beetles and collected the beetles to assess community attributes. We compared the potential future contribution of howler monkeys and other frugivores to seed dispersal based on the seed sizes they disperse in other areas where they occur. In 2 years, howler monkeys consumed 60 animal-dispersed plant species out of the 330 estimated. Twenty-one dung beetle species were attracted to experimentally provided dung; most of them were tunnelers, nocturnal, and large-sized (>10 mm). On average 30% (range 0–100%) of the large seed mimics (14 mm) were moved by dung beetles. About 91% of the species consumed by howlers (size range 0.3–34.3 mm) overlapped in seed size with those removed by dung beetles. In our study area, howler monkeys may consume more large-seeded fruit species than most other frugivores, highlighting their potential to affect forest regeneration. Our results show reintroductions may effectively restore ecological links and enhance ecological processes.     

Forest Fragmentation and Bird Extinctions: San Antonio Eighty Years Later

We report on the extent of bird extinctions at San Antonio, a fragmented cloud forest site in the western Andes of Colombia, for which surveys dating back to 1911 and 1959 an available. In 1911, 128 forest bird species were present in San Antonio. Twenty-four species had disappeared by 1959, and since then 16 more species have gone locally extinct, for a total of 40 species or 31% of the original avifauna. We analyzed patterns of extinction in relation to geographic distribution and foraging guilds. We found that in this montane assemblage, being at the limits of the altitudinal distribution was the main correlate of extinction; 37% of the extinct species were at the upper limit of their altiudinal distribution. We also found that the most vulnerable guilds were the understory insectivores and the large canopy frugivores. Our study illustrates the extent of bird extinctions that are currently undocumented in the highly fragmented forests of the northern Andes, where the absence of baseline information on the fauna of unaltered forests is a limiting factor for the development of conservation and management plans. We stress the need to establish data bases and long-term monitoring projects for the Andean fauna.