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.     

Meta-analysis of the differential effects of habitat fragmentation and degradation on plant genetic diversity

Genetic diversity is a key factor for population survival and evolution. However, anthropogenic habitat disturbance can erode it, making populations more prone to extinction. Aiming to assess the global effects of habitat disturbance on plant genetic variation, we conducted a meta-analysis based on 92 case studies obtained from published literature. We compared the effects of habitat fragmentation and degradation on plant allelic richness and gene diversity (equivalent to expected heterozygosity) and tested whether such changes are sensitive to different life-forms, life spans, mating systems, and commonness. Anthropogenic disturbance had a negative effect on allelic richness, but not on gene diversity. Habitat fragmentation had a negative effect on genetic variation, whereas habitat degradation had no effect. When we examined the individual effects in fragmented habitats, allelic richness and gene diversity decreased, but this decrease was strongly dependent on certain plant traits. Specifically, common long-lived trees and self-incompatible species were more susceptible to allelic richness loss. Conversely, gene diversity decreased in common short-lived species (herbs) with self-compatible reproduction. In a wider geographical context, tropical plant communities were more sensitive to allelic richness loss, whereas temperate plant communities were more sensitive to gene diversity loss. Our synthesis showed complex responses to habitat disturbance among plant species. In many cases, the absence of effects could be the result of the time elapsed since the disturbance event or reproductive systems favoring self-pollination, but attention must be paid to those plant species that are more susceptible to losing genetic diversity, and appropriate conservation should be actions taken.     

Incorporating Climate Science in Applications of the U.S. Endangered Species Act for Aquatic Species

Aquatic species are threatened by climate change but have received comparatively less attention than terrestrial species. We gleaned key strategies for scientists and managers seeking to address climate change in aquatic conservation planning from the literature and existing knowledge. We address 3 categories of conservation effort that rely on scientific analysis and have particular application under the U.S. Endangered Species Act (ESA): assessment of overall risk to a species; long‐term recovery planning; and evaluation of effects of specific actions or perturbations. Fewer data are available for aquatic species to support these analyses, and climate effects on aquatic systems are poorly characterized. Thus, we recommend scientists conducting analyses supporting ESA decisions develop a conceptual model that links climate, habitat, ecosystem, and species response to changing conditions and use this model to organize analyses and future research. We recommend that current climate conditions are not appropriate for projections used in ESA analyses and that long‐term projections of climate‐change effects provide temporal context as a species‐wide assessment provides spatial context. In these projections, climate change should not be discounted solely because the magnitude of projected change at a particular time is uncertain when directionality of climate change is clear. Identifying likely future habitat at the species scale will indicate key refuges and potential range shifts. However, the risks and benefits associated with errors in modeling future habitat are not equivalent. The ESA offers mechanisms for increasing the overall resilience and resistance of species to climate changes, including establishing recovery goals requiring increased genetic and phenotypic diversity, specifying critical habitat in areas not currently occupied but likely to become important, and using adaptive management. Incorporación de las Ciencias Climáticas en las Aplicaciones del Acta Estadunidense de Especies en Peligro para Especies Acuáticas     

Decoupling fragmentation from habitat loss for spiders in patchy agricultural landscapes

Habitat loss reduces species diversity, but the effect of habitat fragmentation on number of species is less clear because fragmentation generally accompanies loss of habitat. We compared four methods that aim to decouple the effects of fragmentation from the effects of habitat loss. Two methods are based on species-area relations, one on Fisher's alpha index of diversity, and one on plots of cumulative number of species detected against cumulative area sampled. We used these methods to analyze the species diversity of spiders in 2, 3.2 × 4 km agricultural landscapes in Southern Judea Lowlands, Israel. Spider diversity increased as fragmentation increased with all four methods, probably not because of the additive within-patch processes, such as edge effect and heterogeneity. The positive relation between fragmentation and species diversity might reflect that most species can disperse through the fields during the wheat-growing season. We suggest that if a given area was designated for the conservation of spiders in Southern Judea Lowlands, Israel, a set of several small patches may maximize species diversity over time.     

Meta‐Analysis of the Effects of Forest Fragmentation on Interspecific Interactions

Forest fragmentation dramatically alters species persistence and distribution and affects many ecological interactions among species. Recent studies suggest that mutualisms, such as pollination and seed dispersal, are more sensitive to the negative effects of forest fragmentation than antagonisms, such as predation or herbivory. We applied meta‐analytical techniques to evaluate this hypothesis and quantified the relative contributions of different components of the fragmentation process (decreases in fragment size, edge effects, increased isolation, and habitat degradation) to the overall effect. The effects of fragmentation on mutualisms were primarily driven by habitat degradation, edge effects, and fragment isolation, and, as predicted, they were consistently more negative on mutualisms than on antagonisms. For the most studied interaction type, seed dispersal, only certain components of fragmentation had significant (edge effects) or marginally significant (fragment size) effects. Seed size modulated the effect of fragmentation: species with large seeds showed stronger negative impacts of fragmentation via reduced dispersal rates. Our results reveal that different components of the habitat fragmentation process have varying impacts on key mutualisms. We also conclude that antagonistic interactions have been understudied in fragmented landscapes, most of the research has concentrated on particular types of mutualistic interactions such as seed dispersal, and that available studies of interspecific interactions have a strong geographical bias (arising mostly from studies carried out in Brazil, Chile, and the United States). Meta‐Análisis de los Efectos de la Fragmentación del Bosque sobre las Interacciones Interespecíficas     

Effects of Growth Form and Functional Traits on Response of Woody Plants to Clearing and Fragmentation of Subtropical Rainforest

The conservation implications of large‐scale rainforest clearing and fragmentation on the persistence of functional and taxonomic diversity remain poorly understood. If traits represent adaptive strategies of plant species to particular circumstances, the expectation is that the effect of forest clearing and fragmentation will be affected by species functional traits, particularly those related to dispersal. We used species occurrence data for woody plants in 46 rainforest patches across 75,000 ha largely cleared of forest by the early 1900s to determine the combined effects of area reduction, fragmentation, and patch size on the taxonomic structure and functional diversity of subtropical rainforest. We compiled species trait values for leaf area, seed dry mass, wood density, and maximum height and calculated species niche breadths. Taxonomic structure, trait values (means, ranges), and the functional diversity of assemblages of climbing and free‐standing plants in remnant patches were quantified. Larger rainforest patches had higher species richness. Species in smaller patches were taxonomically less related than species in larger patches. Free‐standing plants had a high percentage of frugivore dispersed seeds; climbers had a high proportion of small wind‐dispersed seeds. Connections between the patchy spatial distribution of free‐standing species, larger seed sizes, and dispersal syndrome were weak. Assemblages of free‐standing plants in patches showed more taxonomic and spatial structuring than climbing plants. Smaller isolated patches retained relatively high functional diversity and similar taxonomic structure to larger tracts of forest despite lower species richness. The response of woody plants to clearing and fragmentation of subtropical rainforest differed between climbers and slow‐growing mature‐phase forest trees but not between climbers and pioneer trees. Quantifying taxonomic structure and functional diversity provides an improved basis for conservation planning and management by elucidating the effects of forest‐area reduction and fragmentation. Efectos de la Forma de Crecimiento y Atributos Funcionales en la Respuesta de Plantas Leñosas al Desmonte y Fragmentación de Bosque Lluvioso Subtropical     

The Paradox of Forest Fragmentation Genetics

Abstract:  Theory predicts widespread loss of genetic diversity from drift and inbreeding in trees subjected to habitat fragmentation, yet empirical support of this theory is scarce. We argue that population genetics theory may be misapplied in light of ecological realities that, when recognized, require scrutiny of underlying evolutionary assumptions. One ecological reality is that fragment boundaries often do not represent boundaries for mating populations of trees that benefit from long-distance pollination, sometimes abetted by long-distance seed dispersal. Where fragments do not delineate populations, genetic theory of small populations does not apply. Even in spatially isolated populations, where genetic theory may eventually apply, evolutionary arguments assume that samples from fragmented populations represent trees that have had sufficient time to experience drift, inbreeding, and ultimately inbreeding depression, an unwarranted assumption where stands in fragments are living relicts of largely unrelated predisturbance populations. Genetic degradation may not be as important as ecological degradation for many decades following habitat fragmentation.     

Roads,Interrupted Dispersal,and Genetic Diversity in Timber Rattlesnakes

Abstract: Anthropogenic habitat modification often creates barriers to animal movement, transforming formerly contiguous habitat into a patchwork of habitat islands with low connectivity. Roadways are a feature of most landscapes that can act as barriers or filters to migration among local populations. Even small and recently constructed roads can have a significant impact on population genetic structure of some species, but not others. We developed a research approach that combines fine‐scale molecular genetics with behavioral and ecological data to understand the impacts of roads on population structure and connectivity. We used microsatellite markers to characterize genetic variation within and among populations of timber rattlesnakes (Crotalus horridus) occupying communal hibernacula (dens) in regions bisected by roadways. We examined the impact of roads on seasonal migration, genetic diversity, and gene flow among populations. Snakes in hibernacula isolated by roads had significantly lower genetic diversity and higher genetic differentiation than snakes in hibernacula in contiguous habitat. Genetic‐assignment analyses revealed that interruption to seasonal migration was the mechanism underlying these patterns. Our results underscore the sizeable impact of roads on this species, despite their relatively recent construction at our study sites (7 to 10 generations of rattlesnakes), the utility of population genetics for studies of road ecology, and the need for mitigating effects of roads.     

Negative effects of nitrogen deposition on Swiss butterflies

Nitrogen (N) deposition from agriculture and combustion of fossil fuels is a major threat to plant diversity, but its effects on organisms at higher trophic levels are unclear. We investigated how N deposition may affect species richness and abundance (number of individuals per species) in butterflies. We reviewed the peer-reviewed literature on variables used to explain spatial variation in butterfly species richness and found that vegetation variables appeared to be as important as climate and habitat variables in explaining butterfly species richness. It thus seemed likely that increased N deposition could indirectly affect butterfly communities via its influence on plant communities. To test this prediction, we analyzed data from the Swiss biodiversity monitoring program for vascular plants and butterflies in 383 study sites of 1 km² that are evenly distributed throughout Switzerland. The area has a modeled N deposition gradient of 2–44 kg N ha⁻¹ year⁻¹. We used traditional linear models and structural equation models to infer the drivers of the spatial variation in butterfly species richness across Switzerland. High N deposition was consistently linked to low butterfly diversity, suggesting a net loss of butterfly diversity through increased N deposition. We hypothesize that at low elevations, N deposition may contribute to a reduction in butterfly species richness via microclimatic cooling due to increased plant biomass. At higher elevations, negative effects of N deposition on butterfly species richness may also be mediated by reduced plant species richness. In most butterfly species, abundance was negatively related to N deposition, but the strongest negative effects were found for species of conservation concern. We conclude that in addition to factors such as intensified agriculture, habitat fragmentation, and climate change, N deposition is likely to play a key role in negatively affecting butterfly diversity and abundance.     

Estimating Extinction Risk with Metapopulation Models of Large‐Scale Fragmentation

Habitat loss is the principal threat to species. How much habitat remains—and how quickly it is shrinking—are implicitly included in the way the International Union for Conservation of Nature determines a species’ risk of extinction. Many endangered species have habitats that are also fragmented to different extents. Thus, ideally, fragmentation should be quantified in a standard way in risk assessments. Although mapping fragmentation from satellite imagery is easy, efficient techniques for relating maps of remaining habitat to extinction risk are few. Purely spatial metrics from landscape ecology are hard to interpret and do not address extinction directly. Spatially explicit metapopulation models link fragmentation to extinction risk, but standard models work only at small scales. Counterintuitively, these models predict that a species in a large, contiguous habitat will fare worse than one in 2 tiny patches. This occurs because although the species in the large, contiguous habitat has a low probability of extinction, recolonization cannot occur if there are no other patches to provide colonists for a rescue effect. For 4 ecologically comparable bird species of the North Central American highland forests, we devised metapopulation models with area‐weighted self‐colonization terms; this reflected repopulation of a patch from a remnant of individuals that survived an adverse event. Use of this term gives extra weight to a patch in its own rescue effect. Species assigned least risk status were comparable in long‐term extinction risk with those ranked as threatened. This finding suggests that fragmentation has had a substantial negative effect on them that is not accounted for in their Red List category. Estimación del Riesgo de Extinción Mediante Modelos Metapoblacionales de Fragmentación a Gran Escala     

Local and Landscape Correlates of Primate Distribution and Persistence in the Remnant Lowland Rainforests of the Upper Brahmaputra Valley,Northeastern India

Habitat fragmentation affects species distribution and abundance, and drives extinctions. Escalated tropical deforestation and fragmentation have confined many species populations to habitat remnants. How worthwhile is it to invest scarce resources in conserving habitat remnants within densely settled production landscapes? Are these fragments fated to lose species anyway? If not, do other ecological, anthropogenic, and species‐related factors mitigate the effect of fragmentation and offer conservation opportunities? We evaluated, using generalized linear models in an information‐theoretic framework, the effect of local‐ and landscape‐scale factors on the richness, abundance, distribution, and local extinction of 6 primate species in 42 lowland tropical rainforest fragments of the Upper Brahmaputra Valley, northeastern India. On average, the forest fragments lost at least one species in the last 30 years but retained half their original species complement. Species richness declined as proportion of habitat lost increased but was not significantly affected by fragment size and isolation. The occurrence of western hoolock gibbon (Hoolock hoolock) and capped langur (Trachypithecus pileatus) in fragments was inversely related to their isolation and loss of habitat, respectively. Fragment area determined stump‐tailed (Macaca arctoides) and northern pig‐tailed macaque occurrence (Macaca leonina). Assamese macaque (Macaca assamensis) distribution was affected negatively by illegal tree felling, and rhesus macaque (Macaca mulatta) abundance increased as habitat heterogeneity increased. Primate extinction in a fragment was primarily governed by the extent of divergence in its food tree species richness from that in contiguous forests. We suggest the conservation value of these fragments is high because collectively they retained the entire original species pool and individually retained half of it, even a century after fragmentation. Given the extensive habitat and species loss, however, these fragments urgently require protection and active ecological restoration to sustain this rich primate assemblage. Correlaciones Locales y de Paisaje de la Distribución y Persistencia de Primates en los Bosques Lluviosos Remanentes en el Valle del Alto Brahmaputra, Noreste de India     

Carbon Payments and Low‐Cost Conservation

Abstract: A price on carbon is expected to generate demand for carbon offset schemes. This demand could drive investment in tree‐based monocultures that provide higher carbon yields than diverse plantings of native tree and shrub species, which sequester less carbon but provide greater variation in vegetation structure and composition. Economic instruments such as species conservation banking, the creation and trading of credits that represent biological‐diversity values on private land, could close the financial gap between monocultures and more diverse plantings by providing payments to individuals who plant diverse species in locations that contribute to conservation and restoration goals. We studied a highly modified agricultural system in southern Australia that is typical of many temperate agriculture zones globally (i.e., has a high proportion of endangered species, high levels of habitat fragmentation, and presence of non‐native species). We quantified the economic returns from agriculture and from carbon plantings (monoculture and mixed tree and shrubs) under six carbon‐price scenarios. We also identified high‐priority locations for restoration of cleared landscapes with mixed tree and shrub carbon plantings. Depending on the price of carbon, direct annual payments to landowners of AU$7/ha/year to $125/ha/year (US$6–120/ha/year) may be sufficient to augment economic returns from a carbon market and encourage tree plantings that contribute more to the restoration of natural systems and endangered species habitats than monocultures. Thus, areas of high priority for conservation and restoration may be restored relatively cheaply in the presence of a carbon market. Overall, however, less carbon is sequestered by mixed native tree and shrub plantings.     

Understanding Trait‐Dependent Community Disassembly: Dung Beetles,Density Functions,and Forest Fragmentation

Abstract: Anthropogenic disturbances such as fragmentation are rapidly altering biodiversity, yet a lack of attention to species traits and abundance patterns has made the results of most studies difficult to generalize. We determined traits of extinction‐prone species and present a novel strategy for classifying species according to their population‐level response to a gradient of disturbance intensity. We examined the effects of forest fragmentation on dung beetle communities in an archipelago of 33 islands recently created by flooding in Venezuela. Species richness, density, and biomass all declined sharply with decreasing island area and increasing island isolation. Species richness was highly nested, indicating that local extinctions occurred nonrandomly. The most sensitive dung beetle species appeared to require at least 85 ha of forest, more than many large vertebrates. Extinction‐prone species were either large‐bodied, forest specialists, or uncommon. These explanatory variables were unrelated, suggesting at least 3 underlying causes of extirpation. Large species showed high wing loading (body mass/wing area) and a distinct flight strategy that may increase their area requirements. Although forest specificity made most species sensitive to fragmentation, a few persistent habitat generalists dispersed across the matrix. Density functions classified species into 4 response groups on the basis of their change in density with decreasing species richness. Sensitive and persistent species both declined with increasing fragmentation intensity, but persistent species occurred on more islands, which may be due to their higher baseline densities. Compensatory species increased in abundance following the initial loss of sensitive species, but rapidly declined with increasing fragmentation. Supertramp species (widespread habitat generalists) may be poor competitors but strong dispersers; their abundance peaked following the decline of the other 3 groups. Nevertheless, even the least sensitive species were extirpated or rare on the smallest and most isolated islands.     

Effect of Rotational Shepherding on Demographic and Genetic Connectivity of Calcareous Grassland Plants

Response to habitat fragmentation may not be generalized among species, in particular for plant communities with a variety of dispersal traits. Calcareous grasslands are one of the most species‐rich habitats in Central Europe, but abandonment of traditional management has caused a dramatic decline of calcareous grassland species. In the Southern Franconian Alb in Germany, reintroduction of rotational shepherding in previously abandoned grasslands has restored species diversity, and it has been suggested that sheep support seed dispersal among grasslands. We tested the effect of rotational shepherding on demographic and genetic connectivity of calcareous grassland specialist plants and whether the response of plant populations to shepherding was limited to species dispersed by animals (zoochory). Specifically, we tested competing dispersal models and source and focal patch properties to explain landscape connectivity with patch‐occupancy data of 31 species. We fitted the same connectivity models to patch occupancy and nuclear microsatellite data for the herb Dianthus carthusianorum (Carthusian pink). For 27 species, patch connectivity was explained by dispersal by rotational shepherding regardless of adaptations to zoochory, whereas population size (16% species) and patch area (0% species) of source patches were not important predictors of patch occupancy in most species. [Correction made after online publication, February 25, 2014: Population size and patch area percentages were mistakenly inverted, and have now been fixed.] Microsite diversity of focal patches significantly increased the model variance explained by patch occupancy in 90% of the species. For D. carthusianorum, patch connectivity through rotational shepherding explained both patch occupancy and population genetic diversity. Our results suggest shepherding provides dispersal for multiple plant species regardless of their dispersal adaptations and thus offers a useful approach to restore plant diversity in fragmented calcareous grasslands. Efectos del Pastoreo Rotacional sobre la Conectividad Genética y Demográfica de Plantas de Pastizales Calcáreos     

Conservation Implications of Drastic Reductions in the Smallest and Most Isolated Populations of Giant Pandas

Abstract: In conservation biology, understanding the causes of endangerment is a key step to devising effective conservation strategies. We used molecular evidence (coalescent simulations of population changes from microsatellite data) and historical information (habitat and human population changes) to investigate how the most‐isolated populations of giant pandas (Ailuropoda melanoleuca) in the Xiaoxiangling Mountains became highly endangered. These populations experienced a strong, recent demographic reduction (60‐fold), starting approximately 250 years BP. Explosion of the human population and use of non‐native crop species at the peak of the Qing Empire resulted in land‐use changes, deforestation, and habitat fragmentation, which are likely to have led to the drastic reduction of the most‐isolated populations of giant pandas. We predict that demographic, genetic, and environmental factors will lead to extinction of giant pandas in the Xiaoxiangling Mountains in the future if the population remains isolated. Therefore, a targeted conservation action—translocation—has been proposed and is being implemented by the Chinese goverment.     

Multispecies genetic objectives in spatial conservation planning

Growing threats to biodiversity and global alteration of habitats and species distributions make it increasingly necessary to consider evolutionary patterns in conservation decision making. Yet, there is no clear‐cut guidance on how genetic features can be incorporated into conservation‐planning processes, despite multiple molecular markers and several genetic metrics for each marker type to choose from. Genetic patterns differ between species, but the potential tradeoffs among genetic objectives for multiple species in conservation planning are currently understudied. We compared spatial conservation prioritizations derived from 2 metrics of genetic diversity (nucleotide and haplotype diversity) and 2 metrics of genetic isolation (private haplotypes and local genetic differentiation) in mitochondrial DNA of 5 marine species. We compared outcomes of conservation plans based only on habitat representation with plans based on genetic data and habitat representation. Fewer priority areas were selected for conservation plans based solely on habitat representation than on plans that included habitat and genetic data. All 4 genetic metrics selected approximately similar conservation‐priority areas, which is likely a result of prioritizing genetic patterns across a genetically diverse array of species. Largely, our results suggest that multispecies genetic conservation objectives are vital to creating protected‐area networks that appropriately preserve community‐level evolutionary patterns.     

Bridging the gap between ecophysiological and genetic knowledge to assess the adaptive potential of European beech

In this study we aimed to combine knowledge of the ecophysiology and genetics of European beech to assess the potential of this species to adapt to environmental change. Therefore, we performed field and experimental studies on the genetic and ecophysiological functioning of beech. This information was integrated through a coupled genetic–ecophysiological model for individual trees that was parameterized with information derived from our own studies or from the literature. Using the model, we evaluated the adaptive response of beech stands in two ways: firstly, through sensitivity analyses (of initial genetic diversity, pollen dispersal distance, heritability of selected phenotypic traits, and forest management, representing disturbances) and secondly, through the evaluation of the responses of phenotypic traits and their genetic diversity to four management regimes applied to 10 study plots distributed over Western Europe. The model results indicate that the interval between recruitment events strongly affects the rate of adaptive response, because selection is most severe during the early stages of forest development. Forest management regimes largely determine recruitment intervals and thereby the potential for adaptive responses. Forest management regimes also determine the number of mother trees that contribute to the next generation and thereby the genetic variation that is maintained. Consequently, undisturbed forests maintain the largest amount of genetic variation, as recruitment intervals approach the longevity of trees and many mother trees contribute to the next generation. However, undisturbed forests have the slowest adaptive response, for the same reasons.Gene flow through pollen dispersal may compensate for the loss in genetic diversity brought about by selection. The sensitivity analysis showed that the total genetic diversity of a 2 ha stand is not affected by gene flow if the pollen distance distribution is varied from highly left-skewed to almost flat. However, a stand with a prevailing short-distance gene flow has a more pronounced spatial genetic structure than stands with equal short- and long-distance gene flows. The build-up of a spatial genetic structure is also strongly determined by the recruitment interval. Overall, the modelling results indicate that European beech has high adaptive potential to environmental change if recruitment intervals are short and many mother trees contribute to the next generation.The findings have two implications for modelling studies on the impacts of climate change on forests. Firstly: it cannot be taken for granted that parameter values remain constant over a time horizon of even a few generations – this is particularly important for threshold values subject to strong selection, like budburst, frost hardiness, drought tolerance, as used in species area models. Secondly: forest management should be taken into account in future assessments, as management affects the rate of adaptive response and thereby the response on trees and forests to environmental change, and because few forests are unmanaged. We conclude that a coupled ecophysiological and quantitative genetic tree model is a useful tool for such studies.     

Forest Restoration on a Closed Landfill: Rapid Addition of New Species by Bird Dispersal

Urban areas often contain sizeable pockets of degraded land, such as inactive landfills, that could be reclaimed as wildlife habitat and as connecting links to enhance remnant natural areas. In the northeastern U.S., many such lands fail to undergo natural succession to woodland, instead retaining a weedy, herbaceous cover for many years. We hypothesize that seed dispersal is a limiting factor, and that a form of secondary succession could be stimulated by introducing clusters of trees and shrubs to attract avian seed dispersers. As a direct test, we censused a 1.5-ha experimental plantation on the Fresh Kills Landfill (Staten Island, New York) one year after installation, in search of evidence that the plantation was spreading or increasing in diversity. The 17 planted species, many from coastal scrub forests native to this region, were surviving well but contributed almost no seedlings to the area, in part because only 20% of the installed trees or shrubs were reproductive. Of the 1079 woody seedlings found, 95% came from sources outside the plantation; most (71%) were from fleshy-fruited, bird-dispersed plants from nearby woodland fringes. Although the restoration planting itself had not begun to produce seedlings, it did function as a site for attracting dispersers, who enriched the young community with 20 new species. One-fourth of all new recruits were from nine additional wind-dispersed species. Locations with a high ratio of trees to shrubs had proportionately more recruits, indicating that plant size contributed to disperser attraction. The density of new recruits of each species was dependent on distance from the nearest potential seed source. Introducing native species with the capacity to attract avian dispersers may be the key to success of many restoration programs.     

Effects of Dam‐Induced Landscape Fragmentation on Amazonian Ant–Plant Mutualistic Networks

Mutualistic networks are critical to biological diversity maintenance; however, their structures and functionality may be threatened by a swiftly changing world. In the Amazon, the increasing number of dams poses a large threat to biological diversity because they greatly alter and fragment the surrounding landscape. Tight coevolutionary interactions typical of tropical forests, such as the ant–myrmecophyte mutualism, where the myrmecophyte plants provide domatia nesting space to their symbiotic ants, may be jeopardized by the landscape changes caused by dams. We analyzed 31 ant–myrmecophyte mutualistic networks in undisturbed and disturbed sites surrounding Balbina, the largest Central Amazonian dam. We tested how ant–myrmecophyte networks differ among dam‐induced islands, lake edges, and undisturbed forests in terms of species richness, composition, structure, and robustness (number of species remaining in the network after partner extinctions). We also tested how landscape configuration in terms of area, isolation, shape, and neighborhood alters the structure of the ant–myrmecophyte networks on islands. Ant–myrmecophytic networks were highly compartmentalized in undisturbed forests, and the compartments had few strongly connected mutualistic partners. In contrast, networks at lake edges and on islands were not compartmentalized and were negatively affected by island area and isolation in terms of species richness, density, and composition. Habitat loss and fragmentation led to coextinction cascades that contributed to the elimination of entire ant–plant compartments. Furthermore, many myrmecophytic plants in disturbed sites lost their mutualistic ant partners or were colonized by opportunistic, nonspecialized ants. Robustness of ant–myrmecophyte networks on islands was lower than robustness near lake edges and in undisturbed forest and was particularly susceptible to the extinction of plants. Beyond the immediate habitat loss caused by the building of large dams in Amazonia, persistent edge effects and habitat fragmentation associated with dams had large negative effects on animal–plant mutualistic networks. Efectos de la Fragmentación del Paisaje Inducida por Presas sobre Redes Mutualistas Hormiga‐Planta Amazónicas     

Savanna tree density, herbivores, and the herbaceous community: bottom-up vs. top-down effects

Herbivores choose their habitats both to maximize forage intake and to minimize their risk of predation. For African savanna herbivores, the available habitats range in woody cover from open areas with few trees to dense, almost-closed woodlands. This variation in woody cover or density can have a number of consequences for herbaceous species composition, cover, and productivity, as well as for ease of predator detection and avoidance. Here, we consider two alternative possibilities: first, that tree density affects the herbaceous vegetation, with concomitant "bottom-up" effects on herbivore habitat preferences; or, second, that tree density affects predator visibility, mediating "top-down" effects of predators on herbivore habitat preferences. We sampled sites spanning a 10-fold range of tree densities in an Acacia drepanolobium-dominated savanna in Laikipia, Kenya, for variation in (1) herbaceous cover, composition, and species richness; (2) wild and domestic herbivore use; and (3) degree of visibility obstruction by the tree layer. We then used structural equation modeling to consider the potential influences that tree density may have on herbivores and herbaceous community properties. Tree density was associated with substantial variation in herbaceous species composition and richness. Cattle exhibited a fairly uniform use of the landscape, whereas wild herbivores, with the exception of elephants, exhibited a strong preference for areas of low tree density. Model results suggest that this was not a response to variation in herbaceous-community characteristics, but rather a response to the greater visibility associated with more open places. Elephants, in contrast, preferred areas with higher densities of trees, apparently because of greater forage availability. These results suggest that, for all but the largest species, top-down behavioral effects of predator avoidance on herbivores are mediated by tree density. This, in turn, appears to have cascading effects on the herbaceous vegetation. These results shed light on one of the major features of the "landscape of fear" in which African savanna herbivores exist.