Potential Changes in Tree Species Richness and Forest Community Types following Climate Change

Potential changes in tree species richness and forest community types were evaluated for the eastern United States according to five scenarios of future climate change resulting from a doubling of atmospheric carbon dioxide (CO₂). DISTRIB, an empirical model that uses a regression tree analysis approach, was used to generate suitable habitat, or potential future distributions, of 80 common tree species for each scenario. The model assumes that the vegetation and climate are in equilibrium with no barriers to species migration. Combinations of the individual species model outcomes allowed estimates of species richness (from among the 80 species) and forest type (from simple rules) for each of 2100 counties in the eastern United States. Average species richness across all counties may increase slightly with climatic change. This increase tends to be larger as the average temperature of the climate change scenario increases. Dramatic changes in the distribution of potential forest types were modeled. All five scenarios project the extirpation of the spruce–fir forest types from New England. Outputs from only the two least severe scenarios retain aspen–birch, and they are largely reduced. Maple–beech–birch also shows a large reduction in area under all scenarios. By contrast, oak–hickory and oak–pine types were modeled to increase by 34% and 290%, respectively, averaged over the five scenarios. Although many assumptions are made, these modeled outcomes substantially agree with a limited number of predictions from researchers using paleoecological data or other models. Received 12 May 2000; accepted 20 October 2000.     

Effects of Climate Change on the Potential Species Richness of Mesoamerican Forests

The realized species richness of tropical forests cannot yet be reliably mapped at a regional scale due to lack of systematically collected data. An estimate of the potential species richness (PSR), however, can be produced through the use of species distribution modelling. PSR is interpretable as a climatically determined upper limit to observed species richness. We mapped current PSR and future PSR under climate change scenarios for Mesoamerica by combining the spatial distributions of 2000 tree species as predicted by generalized additive models built from herbaria records and climate layers. An explanatory regression tree was used to extract conditional rules describing the relationship between PSR and climate. The results were summarized by country, ecoregion and protected area status in order to investigate current and possible future variability in PSR in the context of regional biodiversity conservation. Length of the dry season was found to be the key determinant of PSR. Protected areas were found to have higher median PSR values than unprotected areas in most of the countries within the study area. Areas with exceptionally high PSR, however, remain unprotected throughout the region. Neither changes in realized species richness nor extinctions will necessarily follow changes in modelled PSR under climate change. However model output suggests that an increase in temperature of around 3°C, combined with a 20 percent decrease in rainfall could lead to a widespread reduction of around 15 percent of current PSR, with values of up to 40 percent in some moist lower montane tropical forests. The modelled PSR of dry forest ecoregions was found to be relatively stable. Some cooler upper montane forests in northern Mesoamerica, where few species of tropical origin are currently found, may gain PSR if species are free to migrate.     

Climate Change Effects on Vegetation Distribution and Carbon Budget in the United States

The Kyoto protocol has focused the attention of the public and policymarkers on the earth's carbon (C) budget. Previous estimates of the impacts of vegetation change have been limited to equilibrium “snapshots” that could not capture nonlinear or threshold effects along the trajectory of change. New models have been designed to complement equilibrium models and simulate vegetation succession through time while estimating variability in the C budget and responses to episodic events such as drought and fire. In addition, a plethora of future climate scenarios has been used to produce a bewildering variety of simulated ecological responses. Our objectives were to use an equilibrium model (Mapped Atmosphere–Plant–Soil system, or MAPSS) and a dynamic model (MC1) to (a) simulate changes in potential equilibrium vegetation distribution under historical conditions and across a wide gradient of future temperature changes to look for consistencies and trends among the many future scenarios, (b) simulate time-dependent changes in vegetation distribution and its associated C pools to illustrate the possible trajectories of vegetation change near the high and low ends of the temperature gradient, and (c) analyze the extent of the US area supporting a negative C balance. Both models agree that a moderate increase in temperature produces an increase in vegetation density and carbon sequestration across most of the US with small changes in vegetation types. Large increases in temperature cause losses of C with large shifts in vegetation types. In the western states, particularly southern California, precipitation and thus vegetation density increase and forests expand under all but the hottest scenarios. In the eastern US, particularly the Southeast, forests expand under the more moderate scenarios but decline under more severe climate scenarios, with catastrophic fires potentially causing rapid vegetation conversions from forest to savanna. Both models show that there is a potential for either positive or negative feedbacks to the atmosphere depending on the level of warming in the climate change scenarios. Received 12 May 2000; accepted 22 November 2000.     

物种丰富度垂直分布格局及影响机制

物种丰富度分布格局是一定地域内物种丰富度沿三维空间的立体分布,包括物种丰富度在经度、纬度和垂直梯度(海拔高度和海水深度)三个维度上的空间分异。近年来物种多样性的垂直分布格局与机制研究得到了生物地理学家和生态学家的重视。物种丰富度的垂直分布格局存在多种类型,但随海拔增加而物种数减少的单调递减模型和中海拔物种丰富度最高的单峰模型较为常见。目前在机制研究中验证较多的是气候稳定性、生物因子(种间相互作用)、能量、生境异质性、干扰、进化时间、物种分化速率、面积、中域效应(mid-domain effect)、生态位保守性(niche conservatism)等假说和机制。物种丰富度的分布格局是多方面因素综合作用的结果;由于地理、地形、气候、地质演化历史、物种库和进化历史、物种分化速率、干扰等差异,在不同地区存在着特别的物种丰富度空间分布格局和机制;处于同一地区的不同类群的物种也因进化扩散历史和生态适应能力不同而呈现多样化的分布格局。因此,对不同地区和类群的物种丰富度格局和机制进行研究应具体分析后才能得到可信结论。     

Fundamental Causes of the Global Patterns of Species Range and Richness1

Global patterns of species range and richness are a consequence of many interacting factors, including environmental conditions, competition, geographical area, and historical/evolutionary development. Two widely studied global patterns of distribution are the latitudinal and elevation gradients of species range and richness. The fundamental mechanisms by which environment and physiology of the plants themselves interact to generate global-scale correlations between increased species range or decreased species richness and latitude/elevation have not previously been established. This paper develops the hypothesis that the primary climatic variables determining global-scale gradients in ectotherm species range and richness are temperature (T) and temperature variability (T), and that the primary physiological variable defining adaptation of ectotherms to temperature is respiratory energy metabolism. This hypothesis is based on a postulate that adaptation of ectotherms to latitudinal/altitudinal gradients of T and T leads to corresponding gradients in properties of energy metabolism. The gradients of metabolic properties give rise to gradients of species range and richness that are observed on a global scale. We demonstrate that natural selection results in ectotherms with metabolic properties matched to their environment and that energy use efficiency and the temperature range allowing growth are inversely related. Thus, opposing selective pressures to increase metabolic energy-use-efficiency or to increase the probability of surviving climate extremes control adaptation of ectotherms to climate. The principles developed in this paper yield fundamental laws of ecology that allow calculation of the contributions of global temperature patterns to the formation of gradients of species range and diversity. Relative values of richness and range are calculated solely from data on abiotic variables. Predictions agree with known patterns of ectotherm distribution.     

Effects of Tree Genotypic Diversity and Species Diversity on the Arthropod Community Associated with Big‐leaf Mahogany

Despite potential interactive effects of plant species and genotypic diversity (SD and GD, respectively) on consumers, studies have usually examined these effects separately. We evaluated the individual and combined effects of tree SD and mahogany (Swietenia macrophylla) GD on the arthropod community associated with mahogany. We conducted this study within the context of a tree diversity experiment consisting of 74 plots with 64 saplings/plot. We sampled 24 of these plots, classified as monocultures of mahogany or polycultures of four species (including mahogany). Within each plot type, mahogany was represented by either one or four maternal families. We surveyed arthropods on mahogany and estimated total arthropod abundance and species richness, as well as abundance and richness separately for herbivorous and predatory arthropods. Overall tree SD and mahogany GD had positive effects on total arthropod species richness and abundance on mahogany, and also exerted interactive effects on total species richness (but not abundance). Analyses conducted by trophic level group showed contrasting patterns; SD positively influenced herbivore species richness but not abundance, and did not affect either predator richness or abundance. GD influenced predator species richness but not abundance, and did not influence herbivore abundance or richness. There were interactive effects of GD and SD only for predator species richness. These results provide evidence that intra‐ and inter‐specific plant diversity exert interactive controls on associated consumer communities, and that the relative importance of SD and GD may vary among higher trophic levels, presumably due to differences in the underlying mechanisms or consumer traits.     

热带森林树种多度和丰富度空间分布特征研究

用巴拿马50 hm2森林动态监测样地内直径≥1 cm的树种资料,分析了该样地树种多度(个体数)和丰富度(物种数)及其方差和变异系数在6个取样尺度(5 m×5 m,10 m×10 m,20 m×20 m,25 m×25 m,50 m×50 m,100 m×100 m)的变化规律.结果显示:(1)由于多度的可加性,不同取样尺度在样地内树种多度的变化表现出一致性;随取样尺度的增加,多度方差呈线性增加,而变异系数呈线性减小.(2)丰富度随取样尺度的变化较为复杂,随取样尺度的增加,丰富度方差呈非线性变化,在取样尺度为25 m×25 m时方差最大;变异系数随取样尺度的增加而呈线性减小.研究表明,大尺度的多度值可以由小尺度的多度值通过外推法估计,而丰富度却不能,在生物多样性的保护和管理中不能简单地从一个取样尺度的生物丰富度推测另一个取样尺度丰富度.     

Effects of Species Richness on Resident and Target Species Components in a Prairie Restoration

The ecological role of biodiversity in achieving successful restoration has been little explored in restoration ecology. We tested the prediction that we are more likely to create persistent, species‐rich plant communities by increasing the number of species sown, and, to some degree, by varying functional group representation, in experimental prairie plantings. There were 12 treatments consisting of 1‐, 2‐, 3‐, 4‐, 8‐, 12‐, and 16‐species mixtures of native perennials representing four functional groups (C₄ grasses, C₃ grasses, nitrogen‐fixing species, and late‐flowering composites) that predominate within Central Plains tallgrass prairies. In 2000, species were seeded into square plots (6 × 6 m), with five replicates per treatment, on former agricultural land. Annually, we measured total species richness and evenness, target species richness and cover, and richness and cover of resident species (i.e., those emerging from the seed bank). Both target species richness and rate of establishment of target communities were highest in the most species‐rich mixtures, but there was no additional benefit for treatments that contained more than eight species. Richness of resident species did not vary with target species richness; however, cover by resident species was lower in the higher target species treatments. Our results, indicating that establishment of species‐rich prairie mimics can be enhanced by starting with larger numbers of species at the outset, have implications for grassland restoration in which community biodiversity creation and maintenance are key goals.     

The Relative Impacts of Climate and Land-Use Change on Conterminous United States Bird Species from 2001 to 2075

Species distribution models often use climate data to assess contemporary and/or future ranges for animal or plant species. Land use and land cover (LULC) data are important predictor variables for determining species range, yet are rarely used when modeling future distributions. In this study, maximum entropy modeling was used to construct species distribution maps for 50 North American bird species to determine relative contributions of climate and LULC for contemporary (2001) and future (2075) time periods. Species presence data were used as a dependent variable, while climate, LULC, and topographic data were used as predictor variables. Results varied by species, but in general, measures of model fit for 2001 indicated significantly poorer fit when either climate or LULC data were excluded from model simulations. Climate covariates provided a higher contribution to 2001 model results than did LULC variables, although both categories of variables strongly contributed. The area deemed to be “suitable” for 2001 species presence was strongly affected by the choice of model covariates, with significantly larger ranges predicted when LULC was excluded as a covariate. Changes in species ranges for 2075 indicate much larger overall range changes due to projected climate change than due to projected LULC change. However, the choice of study area impacted results for both current and projected model applications, with truncation of actual species ranges resulting in lower model fit scores and increased difficulty in interpreting covariate impacts on species range. Results indicate species-specific response to climate and LULC variables; however, both climate and LULC variables clearly are important for modeling both contemporary and potential future species ranges.     

Potential Changes in the Distributions of Western North America Tree and Shrub Taxa under Future Climate Scenarios

Increases in atmospheric greenhouse gases are driving significant changes in global climate. To project potential vegetation response to future climate change, this study uses response surfaces to describe the relationship between bioclimatic variables and the distribution of tree and shrub taxa in western North America. The response surfaces illustrate the probability of the occurrence of a taxon at particular points in climate space. Climate space was defined using three bioclimatic variables: mean temperature of the coldest month, growing degree days, and a moisture index. Species distributions were simulated under present climate using observed data (1951–80, 30-year mean) and under future climate (2090–99, 10-year mean) using scenarios generated by three general circulation models—HADCM2, CGCM1, and CSIRO. The scenarios assume a 1% per year compound increase in greenhouse gases and changes in sulfate (SO₄) aerosols based on the Intergovernmental Panel on Climate Change (IPCC) IS92a scenario. The results indicate that under future climate conditions, potential range changes could be large for many tree and shrub taxa. Shifts in the potential ranges of species are simulated to occur not only northward but in all directions, including southward of the existing ranges of certain species. The simulated potential distributions of some species become increasingly fragmented under the future climate scenarios, while the simulated potential distributions of other species expand. The magnitudes of the simulated range changes imply significant impacts to ecosystems and shifts in patterns of species diversity in western North America. Received 12 May 2000; accepted 20 December 2000.     

When Invasive Plants Disappear: Transformative Restoration Possibilities in the Western United States Resulting from Climate Change

Most ecologists believe that climate change poses a significant threat to the persistence of native species. However, in some areas climate change may reduce or eliminate non-native invasive species, creating opportunities for restoration. If invasive species are no longer suited to novel climate conditions, the native communities that they replaced may not be viable either. If neither invasive nor native species are climatically viable, a type of "transformative" restoration will be required, involving the translocation of novel species that can survive and reproduce under new climate conditions. Here, we illustrate one approach for restoration planning by using bioclimatic envelope modeling to identify restoration opportunities in the western United States, where the invasive plant cheatgrass ( Bromus tectorum ) is no longer climatically viable under 2100 conditions projected by the Geophysical Fluid Dynamics Laboratory (GFDL2.1) coupled atmosphere-ocean general circulation model. We then select one example of a restoration target area and identify novel plant species that could become viable at the site in the wake of climate change. We do so by identifying the closest sites that currently have climate conditions similar to those projected at the restoration target area in 2100. This approach is a first step toward identifying appropriate species for transformative restoration.     

Pond-Breeding Amphibian Species Richness and Habitat Selection in a Beaver-Modified Landscape

ABSTRACT Beaver (Castor canadensis) activity creates wetland habitats with varying hydroperiods important in maintaining habitat diversity for pond-breeding amphibians with significantly different breeding habitat requirements. We documented pond-breeding amphibian assemblages in 71 freshwater wetlands in Acadia National Park, Maine, USA. Using 15 variables describing local pond conditions and wetland landscape characteristics, we developed a priori models to predict sites with high amphibian species richness and used model selection with Akaike's Information Criterion to judge the strength of evidence supporting each model. We developed single-species models to predict wood frog (Rana sylvatica), bullfrog (R. catesbeiana), and pickerel frog (R. palustris) breeding site selection. Sites with high species richness were best predicted by 1) connectivity of wetlands in the landscape through stream corridors and 2) wetland modification by beaver. Wood frog breeding habitat was best predicted by temporary hydroperiod, lack of fish, and absence of current beaver activity. Wood frog breeding was present in abandoned beaver wetlands nearly as often as in nonbeaver wetlands. Bullfrog breeding was limited to active beaver wetlands with fish and permanent water. Pickerel frog breeding sites were best predicted by connectivity through stream corridors within the landscape. As beavers have recolonized areas of their former range in North America, they have increased the number and diversity of available breeding sites in the landscape for pond-breeding amphibians. The resulting mosaic of active and abandoned beaver wetlands both supports rich amphibian assemblages and provides suitable breeding habitat for species with differing habitat requirements. Land managers should consider the potential benefits of minimal management of beavers in promoting and conserving amphibian and wetland diversity at a landscape scale.     

Predicting the Potential Future Distribution of Four Tree Species in Ohio Using Current Habitat Availability and Climatic Forcing

Weinvestigated the effect of habitat loss on the ability of trees to shift in distribution across a landscape dominated by agriculture. The potential distribution shifts of four tree species (Diospyros virginiana, Oxydendron arboreum, Pinus virginiana, Quercus falcata var. falcata) whose northern distribution limits fall in the southern third of Ohio were used to assess possible distribution shift scenarios as a result of global warming. Our predictions derive from the results of simulations using (a) forest inventory based estimates of current distribution and abundance of target species; (b) a satellite-based estimate of forest habitat availability; and (c) a tree migration model (SHIFT). The current distribution and abundance of trees was estimated using USDA Forest Service's Forest Inventory Analysis data and distribution maps from the late 1960s; pre-European settlement forest–nonforest maps were used to represent the fully forested condition for calibration and comparison. Habitat-availability estimates in Ohio were estimated using classified Landsat Thematic Mapper (TM) data from 1994. Tree abundance, forest availability and migration were modeled using a 1-km² pixel size. Forest availability was estimated as the proportion of forested TM pixels within each cell. The probability of a migrating species colonizing an unoccupied cell is modeled as a function of forest availability and distance to occupied cells. The results of the migration models suggest that the species studied are capable of colonizing virtually any forested location within Ohio over the next 100 years if climatic controls over the current distribution that may currently inhibit northward movement are relaxed. The contiguous distribution of these species, however, is not likely to shift more than 10 km during the next century regardless of the magnitude of the climate change. Examining the sensitivity of our simulations by varying critical model attributes, we found that whereas the variables controlling the amount of long-distance dispersal have strong effects on migration rates in the fully forested 1800 situation, they have significantly lesser effects on projections of future migration into highly fragmented forests. The low forest availability that characterizes much of the current Ohio landscape, along with the low likelihood of long distance dispersal, result in potential distribution shifts that are concentrated within the principally forested corridors in southeastern Ohio. We propose that in contrast to the past, future tree migrations are likely to be spatially and temporally correlated as a result of large climatic forcing and channelization through limited regions of available habitat. With respect to the management of biodiversity, this result suggests that it may be very difficult to discern plant migrations of native forest species owing to exceedingly slow rates of movement. Received 19 September 2000; Accepted 2 March 2001.     

Effects of Unseeded Areas on Species Richness of Coal Mines Reclaimed with Municipal Biosolids

Land application of municipal biosolids on coal mine spoils can benefit vegetation establishment in mine reclamation. However, the application of biosolids leads to domination by early‐successional species, such as grasses, and low establishment of woody and volunteer species, thus reducing potential for forestry as a postmining land use. In this experiment, tree seedlings were planted in strips (0.6‐, 1‐, and 4‐m wide) that were not seeded with grasses, and the effects of unseeded strip width on seedling growth and species richness were assessed. Planted seedling mortality was high; therefore, the effect of unseeded strip width on seedling growth could not be determined. However, it was found that natural plant invasion and species richness were highest in the 4‐m unseeded strips. The practice of leaving 4‐m‐wide unseeded strips in mine reclamation with biosolids in the eastern United States, along with the improvement of tree seedling planting practices and planting stock, would help promote a more species‐rich plant community that could be utilized for forestry or a variety of other postmining land uses.     

The influence of past and present climate on the biogeography of modern mammal diversity

Within most terrestrial groups of animals, including mammals, species richness varies along two axes of environmental variation, representing energy availability and plant productivity. This relationship has led to a search for mechanistic links between climate and diversity. Explanations have traditionally focused on single mechanisms, such as variation in environmental carrying capacity or evolutionary rates. Consensus, though, has proved difficult to achieve and there is growing appreciation that geographical patterns of species richness are a product of many interacting factors including biogeographic history and biological traits. Here, we review some current hypotheses on the causes of gradients in mammal richness and range sizes since the two quantities are intimately linked. We then present novel analyses using recent datasets to explore the structure of the environment-richness relationship for mammals. Specifically, we consider the impact of glaciation on present day mammalian diversity gradients. We conclude that not only are multiple processes important in structuring diversity gradients, but also that different processes predominate in different places.