Changes in Species Abundance, Distribution, and Diversity in a Central European Bird Community

We analyzed possible causes of changes in species abundance, range size, and diversity as well as extinctions and colonizations in a central European bird community. Using data from the semiquantitative "Lake Constance" breeding bird atlas, we demonstrated that changes in regional abundances from 1980–1981 to 1990–1992 of 151 coexisting bird species were influenced by breeding habitat and migratory status. Significant declines were found in populations of farmland species and long-distance migrants. Farmland species lost parts of their ranges but hardly changed in local abundance in sites where they still occurred. In contrast, declines in long-distance migrants were caused by significant declines in local abundance with only slight loss of occupied range. Regional extinctions and colonizations were predictable from overall population trends. For example, all species that went extinct were either farmland species or long-distance migrants. Avian community composition was influenced by disproportionate declines of abundant species. This led to declines in the total number of breeding pairs and in community biomass and to increases in community evenness, but to only slight declines in species richness. Future conservation efforts in Europe need to focus more on farmland species and on understanding causes for the declines of long-distance migrants.     

Demographic Responses by Birds to Forest Fragmentation

Abstract:  Despite intensive recent research on the effects of habitat loss and fragmentation on bird populations, our understanding of underlying demographic causes of population declines is limited. We reviewed avian demography in relation to habitat fragmentation. Then, through a meta-analysis, we compared specific demographic responses by forest birds to habitat fragmentation, providing a general perspective of factors that make some species and populations more vulnerable to fragmentation than others. We obtained data from the scientific literature on dispersal, survival, fecundity, and nesting success of birds. Birds were divided into subgroups on the basis of region, nest site, biogeographical history, and migration strategy. Species most sensitive to fragmentation were ground- or open-nesters nesting in shrubs or trees. Residents were equally sensitive to fragmentation in the Nearctic and Palearctic regions, but Nearctic migrants were more sensitive than Palearctic migrants. Old World species were less sensitive than New World species, which was predicted based on the history of forest fragmentation on these two continents. Pairing success was the variable most associated with fragmentation, suggesting an important role of dispersal. Fledgling number or condition, timing of nesting, and clutch size were not associated with sensitivity to fragmentation, suggesting that negative fragmentation effects on birds do not generally result from diminished food resources with increasing level of fragmentation. Future studies on demographic responses of birds to habitat fragmentation would be more effective if based on a combination of measures that can distinguish among the demographic mechanisms underlying population changes related to habitat fragmentation.     

Demographic Limitations of the Ability of Habitat Restoration to Rescue Declining Populations

Abstract:  Habitat restoration is often recommended in conservation without first evaluating whether populations are in fact habitat limited and thus whether declining populations can be stabilized or recovered through habitat restoration. We used a spatially structured demographic model coupled with a dynamic neutral landscape model to evaluate whether habitat restoration could rescue populations of several generic migratory songbirds that differed in their sensitivity to habitat fragmentation (i.e., severity of edge effects on nesting success). Simulating a best-case scenario, landscapes were instantly restored to 100% habitat before, at, or after habitat loss exceeded the species' vulnerability threshold. The vulnerability threshold is a measure of extinction risk, in which the change in population growth rate ( δλ ) scaled to the rate of habitat loss ( δh ) falls below −1% ( δλ/δh ≤ −0.01). Habitat restoration was most effective for species with low-to-moderate edge sensitivities and in landscapes that had not previously experienced extensive fragmentation. To stabilize populations of species that were highly edge sensitive or any species in heavily fragmented landscapes, restoration needed to be initiated long before the vulnerability threshold was reached. In practice, habitat restoration is generally not initiated until a population is at risk of extinction, but our model results demonstrate that some populations cannot be recovered at this point through habitat restoration alone. At this stage, habitat loss and fragmentation have seriously eroded the species' demographic potential such that halting population declines is limited more by demographic factors than the amount of available habitat. Evidence that populations decline in response to habitat loss is thus not sufficient to conclude that habitat restoration will be sufficient to rescue declining populations.     

Use of data on avian demographics and site persistence during overwintering to assess quality of restored riparian habitat

Monitoring responses by birds to restoration of riparian vegetation is relatively cost-effective, but in most assessments species-specific abundances, not demography, are monitored. Data on birds collected during the nonbreeding season are particularly lacking. We captured birds in mist nets and resighted banded birds to estimate species richness and diversity, abundance, demographic indexes, and site-level persistence of permanent-resident and overwintering migrants in remnant and restored riparian sites in California. Species richness in riparian remnants was significantly higher than in restored sites because abundances of uncommon permanent residents were greater in remnants. Species richness of overwintering migrants did not differ between remnants and restored sites. Responses among overwintering migrants (but not permanent residents) to remnant and restored riparian sites differed. Capture rates were higher in remnant or restored riparian sites for 7 of 10 overwintering migratory species. For Lincoln's Sparrows (Melospiza lincolnii) and White-crowned Sparrows (Zonotrichia leucophrys) proportions of older birds were significantly higher in remnants, even though capture rates of these species were higher in restored sites. Overwinter persistence of 4 migrant species was significantly higher in remnant than in restored sites. A higher proportion of Hermit Thrushes (Catharus guttatus, 56.3%), older Fox Sparrows (Passerella iliaca, 57.1%), Lincoln's Sparrows (59.7%), and White-crowned Sparrows (67.8%) persisted in remnants than restored sites. Our results suggest restored riparian sites provide habitat for a wide variety of species in comparable abundances and diversity as occurs in remnant riparian sites. Our demographic and persistence data showed that remnants supported some species and age classes to a greater extent than restored sites.     

Spatial scaling of avian population dynamics: population abundance, growth rate, and variability

Synchrony in population fluctuations has been identified as an important component of population dynamics. In a previous study, we determined that local-scale (<15-km) spatial synchrony of bird populations in New England was correlated with synchronous fluctuations in lepidopteran larvae abundance and with the North Atlantic Oscillation. Here we address five questions that extend the scope of our earlier study using North American Breeding Bird Survey data. First, do bird populations in eastern North America exhibit spatial synchrony in abundances at scales beyond those we have documented previously? Second, does spatial synchrony depend on what population metric is analyzed (e.g., abundance, growth rate, or variability)? Third, is there geographic concordance in where species exhibit synchrony? Fourth, for those species that exhibit significant geographic concordance, are there landscape and habitat variables that contribute to the observed patterns? Fifth, is spatial synchrony affected by a species' life history traits? Significant spatial synchrony was common and its magnitude was dependent on the population metric analyzed. Twenty-four of 29 species examined exhibited significant synchrony in population abundance: mean local autocorrelation (rho)= 0.15; mean spatial extent (mean distance where rho=0) = 420.7 km. Five of the 29 species exhibited significant synchrony in annual population growth rate (mean local autocorrelation = 0.06, mean distance = 457.8 km). Ten of the 29 species exhibited significant synchrony in population abundance variability (mean local autocorrelation = 0.49, mean distance = 413.8 km). Analyses of landscape structure indicated that habitat variables were infrequent contributors to spatial synchrony. Likewise, we detected no effects of life history traits on synchrony in population abundance or growth rate. However, short-distance migrants exhibited more spatially extensive synchrony in population variability than either year-round residents or long-distance migrants. The dissimilarity of the spatial extent of synchrony across species suggests that most populations are not regulated at similar spatial scales. The spatial scale of the population synchrony patterns we describe is likely larger than the actual scale of population regulation, and in turn, the scale of population regulation is undoubtedly larger than the scale of individual ecological requirements.     

Poleward shifts in winter ranges of North American birds

Climate change is thought to promote the poleward movement of geographic ranges; however, the spatial dynamics, mechanisms, and regional anthropogenic drivers associated with these trends have not been fully explored. We estimated changes in latitude of northern range boundaries, center of occurrence, and center of abundance for 254 species of winter avifauna in North America from 1975 to 2004. After accounting for the effect of range size and the location of the northern boundary, positive latitudinal trends were evident for the northern boundary (1.48 km/yr), center of occurrence (0.45 km/yr), and center of abundance (1.03 km/yr). The northern boundary, when examined across individual species, had the most variable trends (SD = 7.46 km/yr) relative to the center of occurrence (SD = 2.36 km/yr) and center of abundance (SD = 5.57 km/yr). Trends did not differ based on migratory status, but there was evidence that trends differed for species with ranges centered in the southern vs. northern portion of the study area. Species occurred more sporadically over time at northern range boundaries, and northern boundaries were associated with a concentration of colonization and extirpation events, with a greater prevalence of colonization events likely promoting poleward trends. Regional anthropogenic drivers explained approximately 8% of the trend for the northern boundary, 14% for the center of occurrence, and 18% for the center of abundance; however, these effects were localized in the northern portion of species' ranges and were associated with distributional changes within ranges, primarily abundance, producing patterns that mimicked poleward movements. We conclude that poleward distributional shifts represent the interaction between climate change and regional factors whose outcome is determined by the scale of the analysis and the biotic and abiotic features in the region, and how anthropogenic activities have impacted these features.     

Abundance declines of a native forb have nonlinear impacts on grassland invasion resistance

The effects of declining plant biodiversity on ecosystem processes are well studied, with most investigations examining the role of species richness declines rather than declines of species abundance. Using grassland mesocosms, we examined how the abundance of a native, resident species, Hemizonia congesta (hayfield tarweed), affected exotic Centaurea solstitialis (yellow starthistle) invasion. We found that progressive H. congesta abundance declines had threshold effects on invasion resistance, with initial declines resulting in minor increases in invasion and subsequent declines leading to accelerating increases in invader performance. Reduced invasion resistance was explained by increased resource availability as H. congesta declined. We also found evidence that resident abundance might indirectly affect invasion by mediating invader impact on resident competitors; C. solstitialis disproportionately reduced H. congesta biomass in low-abundance rather than high-abundance populations. H. congesta's direct and indirect effects on invasion resistance illustrate that an individual species' declining abundance can have accelerating, deleterious effects on ecosystem functions of conservation value.     

Mitochondrial DNA Variability of the Gray Wolf: Genetic Consequences of Population Decline and Habitat Fragmentation

The gray wolf is a large, highly mobile predator whose original geographic range included most of the Northern Hemisphere. High rates of genetic exchange probably characterized even distantly-separated populations in the past, but recent population declines and habitat fragmentation have isolated previously contiguous populations, especially in the Old World. We examine mitochondrial DNA (mtDNA) variability among twenty-six populations of wolves from throughout their geographic range. We find eighteen mtDNA genotypes in gray wolves, seven of these are derived from hybridization with coyotes, four are confined to the New World, six are confined to the Old World and one is shared by both areas. Genetic differentiation among wolf populations is significant but small in magnitude. In the Old World, most localities have a single unique genotype, whereas in the New World several genotypes occur at most localities and three of the five genotypes are nearly ubiquitous. The pattern of genetic differentiation in the gray wolf contrasts with that of another large, highly vagile canid, the coyote, in which genetic differentiation among populations is not significant even among widely separated localities. We suggest that the difference between these two species reflects the rapid, recent increase in coyote numbers and expansion of their geographic range, and the coincident decline in gray wolf populations. Apparent genetic differences among extant wolf populations may be a recent phenomenon reflecting population declines and habitat fragmentation rather than a long history of genetic isolation.     

A Mechanistic Approach to Evaluation of Umbrella Species as Conservation Surrogates

Abstract:  Although species with large area requirements are sometimes used as umbrella species, their general utility as conservation tools is uncertain. We surveyed the species diversity of birds, butterflies, carabids, and forest-floor plants in forest sites across an area (1600 km²) in which we delineated large breeding home ranges of Northern Goshawk ( Accipiter gentilis ). We tested whether protection of the home ranges could serve as an effective umbrella to protect sympatric species of the four taxa. We also used an empirical habitat model of occupancy of home range to examine mechanisms by which the Northern Goshawk acts as an umbrella species. Among species richness, abundance, and species composition of the four taxa, only abundance and species composition of birds differed between sites located inside and outside home ranges, which was due to greater abundance of bird species that were prey of Northern Goshawks inside the home ranges. Thus, although home range indicated areas with high abundance of certain bird prey species, it was not effective as an indicator of the species diversity of all four taxa. We also did not find any difference in species richness, abundance, and species composition between sites predicted as occupied and unoccupied using the habitat model. In contrast, when we selected sites on the basis of each habitat variable in the model, habitat variables that selected sites either in agricultural or forested landscapes encompassed sites with high species richness or particular species composition. This result suggests that the low performance of the Northern Goshawk as an umbrella species is due to this species' preference for habitat in both agricultural and forested landscapes. Species that can adjust to changes in habitat conditions may not act as effective umbrella species despite having large home ranges.     

Defining Forest Fragmentation by Corridor Width: The Influence of Narrow Forest-Dividing Corridors on Forest-Nesting Birds in Southern New Jersey

In studies of forest fragmentation, a fundamental inconsistency exists in the distance criterion used to define the discreteness of forest fragments. We examined three types of ubiquitous, narrow, forest-dividing corridors for effects that influence the relative abundance and community composition of forest-nesting birds. Fixed-radius (100-meter) point counts were conducted on 54 transects established along three width classes of corridors: unpaved roads (8 meters wide), paved roads (16 meters wide), and powerlines (23 meters wide). Transect locations were distributed equally among corridor edge, forest margin 100 meters from corridor edge, and forest interior 300 meters from corridor edge. Forest-interior species of Neotropical migrants had significantly reduced relative abundances on edge transects along 16- and 23-meter corridors, compared with 8-meter corridors and with forest interior points along all three corridor-width classes. At a landscape scale, the consequences of apparently small reductions in forest area by the presence of narrow forest-dividing corridors may be cumulatively significant for abundances of forest-interior species. Brown-headed Cowbirds were more abundant than 20 of 21 forest-interior Neotropical migrants. We found surprisingly high abundances of cowbirds associated with narrow forest-dividing corridors, especially those with mowed grass. Corridor widths as narrow as 8 meters produce forest fragmentation effects in part by attracting cowbirds and nest predators to corridors and adjacent forest interiors. The most serious implication of this study is that narrow forest-dividing corridors may function as ecological traps for forest-interior Neotropical migrants. We suggest that these widespread corridors may be inconspicuous but important contributors to declines of forest-interior nesting species in eastern North America.     

Reproduction on the edge: large-scale patterns of individual performance in a marine invertebrate

Reproductive output is a central attribute of life history, providing a measure of individual and population performance. The fields of ecology, biogeography, and evolutionary biology take disparate approaches in addressing spatial variation in reproduction, and thus we lack clear predictions for how reproductive output should vary geographically. We empirically investigate these contrasting theoretical approaches by determining geographic patterns in reproductive output for intertidal populations of the purple sea urchin, Strongylocentrotus purpuratus, at 15 sites spanning a large geographic distance (9 degrees span of latitude) from central California, USA, to Baja California, Mexico. Contrary to predictions from biogeography, some of the highest values of reproductive output are at sites near the species' southern range boundary. Additionally, sea urchins do not exhibit a latitudinal gradient in reproduction, but rather show considerable mesoscale variation in reproductive output. Spatial analyses reveal that this variation is correlated with coastal topographic features that are known to influence the pattern of nearshore nutrient flux and circulation. We hypothesize that urchins' reproductive output may be driven by the spatial distribution of their food supply, drift macroalgae, the abundance of which is influenced by both nutrient supply and alongshore transport processes that are coupled to topographic features. Large-scale studies such as this provide valuable insight into the causes of species' range limits, population connectivity, habitat reserve design, and forecasting the effects of climate change on species' distributions.     

Habitat Loss and Changes in the Species-Area Relationship

Abstract: The species-area relationship (SAR) has been used successfully to predict extinction from extent of habitat reduction. These extinction estimates assume that species have uniformly distributed range requirements and a minimum abundance level required for persistence; how many species are lost depends solely on how much habitat is removed, not on where it is removed. We consider another limiting case in which range requirements, rather than abundances, determine extinctions. We used a new method for constructing SARs based on assumptions about geographic ranges of species. Our results show that habitat destruction can change the SAR and consequently the number of species predicted to be lost due to habitat destruction. Our method generates SARs that vary in shape according to the specific distributions of geographic range and occupancy but that have the common feature of being described by a power law with an exponent of <1. When the geographic range of species was included in the SAR, the way habitat was lost became important. Although the SAR before habitat destruction is often used to predict species loss after habitat destruction, assumptions must be clearly stated. To predict the damage caused by habitat loss with our model, it is necessary to know the fraction of aggregated species, the distribution of geographic ranges, the form of habitat destruction, and the sampling protocol. The remaining theoretical challenge is to develop a full theory that links abundance and range.     

The Bias of Complementarity Hotspots toward Marginal Populations

Abstract: It has been suggested that using complementarity to identify networks of important areas for conserving biodiversity may preferentially select areas within the margins of species ranges. We tested this idea by examining the location of complementarity hotspots in relation to two measures of range core-periphery. The first measures patterns of aggregation among records within each species' range to identify areas within the core (i.e., areas with aggregated distributions) and periphery (i.e., areas with scattered distributions) of the range. The second measures spatial turnover among species to identify areas with a high density of range edges. For three selected groups of terrestrial vertebrates in Europe—mammals, birds, and herptiles—areas chosen based on complementarity were located within the margins of species' ranges more often than expected by chance. This pattern was consistent for the two measures of core-periphery we used. The bias of complementarity hotspots toward marginal populations is especially important for species with restricted range sizes. If extinctions are determined mainly by demographic factors, then selecting areas at the peripheries of species' ranges might be a poor option. But if extinctions are determined mainly by extrinsic factors, then peripheral populations might be important to ensure the long-term persistence of species.     

Social dominance in migrant and resident turkey vultures at carcasses: evidence for a despotic distribution?

We placed carcasses in three different vegetation types in the heterogeneous savannas of central Venezuela to investigate the role of social dominance in habitat use by flocking migrant and resident turkey vultures (Cathartes aura meridionalis and C. a. ruficollis). Migrants foraged primarily in savanna habitats while residents foraged almost exclusively in gallery forest. In the gallery forest residents discovered carrion first significantly more often than migrants, despite there being equal densities of residents and migrants foraging over this habitat. Because residents fed in smaller groups than migrants at carcasses they had higher feeding rates. There was also a negative relationship between group sizes of residents and migrants. The feeding rate of residents declined in response to increased group size of migrants, but group size of residents had no effect. Migrant group size also had a greater effect on resident feeding rates than king vulture presence or absence. When the effect of migrant and resident group size on feeding rates in migrants was compared, the most significant factor was migrant group size. A second analysis showed that both resident group size and presence or absence of king vultures had a significant effect on feeding rates in migrants. Rates of agonistic encounters in migrant and resident turkey vultures increased weakly in relation to group size. However, there was an increase in residents' encounter rate with migrants in relation to increased migrant group size; there was no difference in resident encounter rates with other residents in relation to resident group size. Migrants dominated residents in almost all agonistic interactions over carcasses. We suggest that savanna habitats were less attractive to residents for foraging because they held larger groups of migrants.     

Environmental change and declining resource availability for small-mammal communities in the Great Basin

Changes in climate and land use can impact natural systems across all levels of ecological organization. Most documented and anticipated effects consider species' properties, including phenologies, geographic distributions, and abundances. Responses of higher-level aggregate community or ecosystem properties have not been considered as they are assumed to be relatively stable due to compensatory dynamics and diversity-stability relationships. However, this assumption may not be as fundamental as previously thought. Here we assess stability in the aggregate properties of total abundance, biomass, and energy consumption for small-mammal communities in the Great Basin, using paired historical and modern survey data spanning nearly a century of environmental change. Results show marked declines in each aggregate property independent of spatial scale, elevation, or habitat type, and a reallocation of available biomass and energy favoring diet and habitat generalists. Because aggregate properties directly reflect resource availability, our findings indicate a regionwide decline in resources of 50% over the past century, which may signal a resource crisis. This work illustrates the power of using aggregate properties as indicators of ecological conditions and environmental change at broad spatial and temporal scales.     

The distribution of lianas and their change in abundance in temperate forests over the past 45 years

Lianas (woody vines) are an important and dynamic component of many forests throughout the world, and increases in CO2, mean winter temperature, and forest fragmentation may promote their growth and proliferation in temperate forests. In this study, we used a 45-year data set to test the hypothesis that lianas have increased in abundance and basal area in the interiors of 14 deciduous temperate forests in Wisconsin (USA) since 1959. We also censused woody plants along a gradient from the forest edge to the interior in seven of these forests to test the hypothesis that the abundance of lianas declines significantly with increasing distance from the forest edge. We found that lianas did not increase in abundance within the interiors of temperate forests in Wisconsin over the last 45 years. However, relative and absolute liana abundance decreased sharply with increasing distance from forest edges. Our findings suggest that forest fragmentation, not climate change, may be increasing the abundance of lianas in northern deciduous temperate forests, and that lianas may further increase in abundance if the severity of forest fragmentation intensifies.     

Demography of central and marginal populations of monkeyflowers (Mimulus cardinalis and M. lewisii)

Every species occupies a limited geographic area, but how spatiotemporal environmental variation affects individual and population fitness to create range limits is not well understood. Because range boundaries arise where, on average, populations are more likely to go extinct than to persist, range limits are an inherently population-level problem for which a demographic framework is useful. In this study, I compare demographic parameters and population dynamics between central and marginal populations of monkeyflowers, Mimulus cardinalis and M. lewisii, along an elevation gradient spanning both species' ranges. Central and marginal populations of both species differed in survival and fecundity. For M. lewisii, these components of fitness were higher in central than in marginal populations, but for M. cardinalis the converse was true. To assess spatiotemporal variation in population dynamics, I used transition matrix models to estimate asymptotic population growth rates (lambda) and found that population growth rates of M. lewisii were highest at the range center and reduced at the range margin. Population growth rates of M. cardinalis were highest at the range margin and greatly reduced at the range center. Life table response analysis decomposed spatiotemporal variation in lambda into contributions from each transition between life stages, finding that transitions from large nonreproductive and reproductive plants to the seed class and stasis in the reproductive class made the largest contributions to spatial differences in lambda. These transitions had only low to moderate sensitivities, indicating that differences in projected population growth rates resulted mainly from observed differences in transition matrix parameters and their underlying vital rates.     

Contrasting effects of habitat loss and fragmentation on coral-associated reef fishes

Disturbance can result in the fragmentation and/or loss of suitable habitat, both of which can have important consequences for survival, species interactions, and resulting patterns of local diversity. However, effects of habitat loss and fragmentation are typically confounded during disturbance events, and previous attempts to determine their relative significance have proved ineffective. Here we experimentally manipulated live coral habitats to examine the potential independent and interactive effects of habitat loss and fragmentation on survival, abundance, and species richness of recruitment-stage, coral-associated reef fishes. Loss of 75% of live coral from experimental reefs resulted in low survival of a coral-associated damselfish and low abundance and richness of other recruits 16 weeks after habitat manipulations. In contrast, fragmentation had positive effects on damselfish survival and resulted in greater abundance and species richness of other recruits. We hypothesize that spacing of habitat through fragmentation weakens competition within and among species. Comparison of effect sizes over the course of the study period revealed that, in the first six weeks following habitat manipulations, the positive effects of fragmentation were at least four times stronger than the effects of habitat loss. This initial positive effect of fragmentation attenuated considerably after 16 weeks, whereas the negative effects of habitat loss increased in strength over time. There was little indication that the amount of habitat influenced the magnitude of the habitat fragmentation effect. Numerous studies have reported dramatic declines in coral reef fish abundance and diversity in response to disturbances that cause the loss and fragmentation of coral habitats. Our results suggest that these declines occur as a result of habitat loss, not habitat fragmentation. Positive fragmentation effects may actually buffer against the negative effects of habitat loss and contribute to the resistance of reef fish populations to declines in coral cover.     

Range Size and Extinction Risk in Forest Birds

Abstract:  Small geographical range size is the single best predictor of threat of extinction in terrestrial species. Knowing how small a species' range has to be before authorities consider it threatened with extinction would allow prediction of a species' risk from continued deforestation and warming climates and provide a baseline for conservation and management strategies aspiring to mitigate these threats. To determine the threshold at which forest-dependent bird species become threatened with extinction, we compared the range sizes of threatened and nonthreatened species. In doing so, we present a simple, repeatable, and practical protocol to quantify range size. We started with species' ranges published in field guides or comparable sources. We then trimmed these ranges, that is, we included only those parts of the ranges that met the species' requirements of elevation and types of forest preferred. Finally, we further trimmed the ranges to the amount of forest cover that remains. This protocol generated an estimate of the remaining suitable range for each species. We compared these range estimates with those from the World Conservation Union Red List. We used the smaller of the two estimates to determine the threshold, 11,000 km², below which birds should be considered threatened. Species considered threatened that have larger ranges than this qualified under other (nonspatial) red list criteria. We identified a suite of species (18) that have not yet qualified as threatened but that have perilously small ranges—about 11% of the nonthreatened birds we analyzed. These birds are likely at risk of extinction and reevaluation of their status is urgently needed.     

Process-based modeling of species' distributions: what limits temperate tree species' range boundaries?

Niche-based models are widely used to understand what environmental factors determine species' distributions, but they do not provide a clear framework to study the processes involved in defining species' ranges. Here we used a process-based model to identify these processes and to assess the potential distribution of 17 North American boreal/temperate tree species. Using input of only climate and soil properties, the model reproduced the 17 species' distributions accurately. Our results allowed us to identify the climatic factors as well as the biological processes involved in limiting species' ranges. The model showed that climatic constraints limit species' distributions mainly through their impact on phenological processes, and secondarily through their impact on drought and frost mortality. The northern limit of species' ranges appears to be caused mainly by the inability to undergo full fruit ripening and/or flowering, while the southern limit is caused by the inability to flower or by frost injury to flowers. These findings about the ecological processes shaping tree species' distribution represent a crucial step toward obtaining a more complete picture of the potential impact of climate on species' ranges.