Information processing in models for density-dependent emigration: A comparison

Density-dependent emigration has been recognized as a fitness enhancing strategy. Yet, especially in the modelling literature there is no consensus about how density-dependent emigration should quantitatively be incorporated into metapopulation models. In this paper we compare the performance of five different dispersal strategies (defined by the functional link between density and emigration probability). Four of these strategies are based on published functional relationships between local population density and emigration probability, one assumes density-independent dispersal. We use individual-based simulations of time-discrete metapopulation dynamics and conduct evolution experiments for a broad range of values for dispersal mortality and environmental stochasticity. For each set of these conditions we analyze the evolution of emigration rates in ‘monoculture experiments’ (with only one type of dispersal strategy used by all individuals in the metapopulation) as well as in selection experiments that allow a pair-wise comparison of the performance of each functional type. We find that a single-parameter ‘asymptotic threshold’ strategy - derived from the marginal value theorem - with a decelerating increase of emigration rate with increasing population density, out-competes any other strategy, i.e. density-independent emigration, a ‘linear threshold’ strategy and a flexible three-parameter strategy. Only when environmental conditions select for extremely high emigration probabilities (close to one), strategies may perform approximately equally. A simple threshold strategy derived for the case of continuous population growth performs even worse than the density-independent strategy. As the functional type of the dispersal function implemented in metapopulation models may severely affect predictions concerning the survival of populations, range expansion, or community changes we clearly recommend to carefully select adequate functions to model density-dependent dispersal.     

Exploring the internal and external wildlife gradients created by conservation fences

Spillover effects are an expansion of conservation benefits beyond protected areas through dispersal of species that reside within. They have been well documented in marine but not terrestrial systems. To understand the effects on wildlife created by conservation fences, we explored the internal and external gradients of activity in mammal, reptile, and bird species at a conservation reserve in arid Australia that is fenced to exclude invasive rabbits (Oryctolagus cuniculus), cats (Felis catus), and foxes (Vulpes vulpes). Two methods were used: counts of animal tracks along transects on sand dunes and captures at pitfall-trapping sites. In both cases, sites were spaced at different distances from the reserve fenceline inside and outside the reserve. We recorded a range of spillover, source-sink, step, and barrier effects that combined to create a zone within and around the reserve with fence-induced species-specific wildlife gradients. Two endemic rodents but none of the 4 mammal species reintroduced to the reserve showed positive spillover effects. Barrier effects, where activity was highest close to the fence, were recorded for the feral cat and native bettong (Bettongia lesueur), species that could not breach the fence. In comparison, some reptiles and native mammal species that could permeate the fence displayed source-sink effects; that is, their activity levels were reduced close to the fence likely due to constant emigration to the side with lower density. Activity of some reptiles was lowest at sites inside the reserve and gradually increased at outside sites with distance from the fence, a gradient likely related to trophic cascades triggered by predator exclusion. Our result shows that fenced reserves can create overlapping layers of species-specific gradients related to each species’ ability to permeate the fence and its varying susceptibility to threats. Managers should be aware that these gradients may extend for several kilometers either side of the fence and that not all contained species will increase in abundance. Creating wider conservation benefits may require increased fence permeability and threat reduction outside the fence.     

Modeling range dynamics in heterogeneous landscapes: invasion of the hemlock woolly adelgid in eastern North America

Range expansion by native and exotic species will continue to be a major component of global change. Anticipating the potential effects of changes in species distributions requires models capable of forecasting population spread across realistic, heterogeneous landscapes and subject to spatiotemporal variability in habitat suitability. Several decades of theory and model development, as well as increased computing power and availability of fine-resolution GIS data, now make such models possible. Still unanswered, however, is the question of how well this new generation of dynamic models will anticipate range expansion. Here we develop a spatially explicit stochastic model that combines dynamic dispersal and population processes with fine-resolution maps characterizing spatiotemporal heterogeneity in climate and habitat to model range expansion of the hemlock woolly adelgid (HWA; Adelges tsugae). We parameterize this model using multiyear data sets describing population and dispersal dynamics of HWA and apply it to eastern North America over a 57-year period (1951-2008). To evaluate the model, the observed pattern of spread of HWA during this same period was compared to model predictions. Our model predicts considerable heterogeneity in the risk of HWA invasion across space and through time, and it suggests that spatiotemporal variation in winter temperature, rather than hemlock abundance, exerts a primary control on the spread of HWA. Although the simulations generally matched the observed current extent of the invasion of HWA and patterns of anisotropic spread, it did not correctly predict when HWA was observed to arrive in different geographic regions. We attribute differences between the modeled and observed dynamics to an inability to capture the timing and direction of long-distance dispersal events that substantially affected the ensuing pattern of spread.     

Are Swiss birds tracking climate change?: Detecting elevational shifts using response curve shapes

Climate change is affecting biodiversity worldwide inducing species to either “move, adapt or die”. In this paper we propose a conceptual framework for analysing range shifts, namely a catalogue of the possible patterns of change in the distribution of a species along elevational or other environmental gradients and an improved quantitative methodology to identify and objectively describe these patterns. Patterns are defined in terms of changes occurring at the leading, trailing or both edges of the distribution: (a) leading edge expansion, (b) trailing edge retraction, (c) range expansion, (d) optimum shift, (e) expansion, (f) retraction, and (g) shift. The methodology is based on the modelling of species distributions along a gradient using generalized additive models (GAMs). Separate models are calibrated for two distinct periods of assessment and response curves are compared over five reference points. Changes occurred at these points are formalized into a code that ultimately designates the corresponding change pattern. We tested the proposed methodology using data from the Swiss national common breeding bird survey. The elevational distributions of 95 bird species were modelled for the periods 1999-2002 and 2004-2007 and significant upward shifts (all patterns confounded) were identified for 35% of the species. Over the same period, an increase in mean temperature was registered for Switzerland. In consideration of the short period covered by the case study, assessed change patterns are considered to correspond to intermediate patterns in an ongoing shifting process. However, similar patterns can be determined by habitat barriers, land use/land cover changes, competition with concurrent or invasive species or different warming rates at different elevations.     

The effect of positive interactions on community structure in a multi-species metacommunity model along an environmental gradient

Positive interactions are widely recognized as playing a major role in the organization of community structure and diversity. As such, recent theoretical and empirical works have revealed the significant contribution of positive interactions in shaping species’ geographical distributions, particularly in harsh abiotic conditions. In this report, we explore the joint influence of local dispersal and an environmental gradient on the spatial distribution, structure and function of communities containing positive interactions. While most previous theoretical efforts were limited to modelling the dynamics of single pairs of associated species being mutualist or competitor, here we employ a spatially explicit multi-species metacommunity model covering a rich range of interspecific interactions (mutualism, competition and exploitation) along an environmental gradient. We find that mutualistic interactions dominate in communities with low diversity characterized by limited species dispersal and poor habitat quality. On the other hand, the fraction of mutualistic interactions decreases at the expense of exploitation and competition with the increase in diversity caused by higher dispersal and/or habitat quality. Our multi-species model exemplifies the ubiquitous presence of mutualistic interactions and the role of mutualistic species as facilitators for the further establishment of species during ecosystem assembly. We therefore argue that mutualism is an essential component driving the origination of complex and diverse communities.     

The Role of Roadsides in Plant Invasions: a Demographic Approach

Abstract:  Non-native plant species are common along roadsides, but presence does not necessarily indicate spread along the road axis. Roadsides may serve merely as habitat for a species spreading independently of roads. The potential conduit function of roads depends on the habitat specificity of the spreading species, its dispersal range relative to the spacing of roads in the landscape, and the relative importance of long- and short-range dispersal. We describe a demographic model of the road × species interaction and suggest methods of assessing conduit function in the field based on the model results. A species limited to roadside habitat will be constrained to spread along the road axis unless its long-range dispersal is sufficient to carry it across the intervening unfavorable area to another road. It will propagate along a road corridor at a rate determined by the scale of short-range dispersal. Effective management of an invasion requires distinguishing between the habitat and conduit functions, a distinction difficult to make with only snapshot data. Invasions can be reconstructed by several methods, but none is totally satisfactory. We suggest comparing stem distributions on transects parallel and perpendicular to the road axis, and beside the road, and away from it, with an idealized Gaussian curve. Such comparisons would allow discrimination between pattern determined by habitat suitability and pattern reflecting random and facilitated dispersal.     

Explaining long-distance dispersal: effects of dispersal distance on survival and growth in a stream salamander

Long-distance dispersal (LDD) may contribute disproportionately to range expansions, the creation of new evolutionary lineages, and species persistence in human-dominated landscapes. However, because data on the individual consequences of dispersal distance are extremely limited, we have little insight on how LDD is maintained in natural populations. I used six years of spatially explicit capture-mark-recapture (CMR) data to test the prediction that individual performance increases with dispersal distance in the stream salamander Gyrinophilus porphyriticus. Dispersal distance was total distance moved along the 1-km study stream, ranging from 0 to 565 m. To quantify individual performance, I used CMR estimates of survival and individual growth rates based on change in body length. Survival and growth rates increased significantly with dispersal distance. These relationships were not confounded by pre-dispersal body condition or by ecological gradients along the stream. Individual benefits of LDD were likely caused by an increase in the upper limit of settlement site quality with dispersal distance. My results do not support the view that the fitness consequences of LDD are unpredictable and instead suggest that consistent evolutionary mechanisms may explain the prevalence of LDD in nature. They also highlight the value of direct CMR data for understanding the individual consequences of variation in dispersal distance and how that variation is maintained in natural populations.     

The control of emigration and its consequences for the survival of populations

Dispersal is the key process enhancing the long-term persistence of metapopulations in heterogeneous and dynamic landscapes. However, any individual emigrating from a occupied patch also increases the risk of local population extinction. The consequences of this increase for metapopulation persistence likely depend on the control of emigration. In this paper, we present results of individual-based simulations to compare the consequences of density-independent (DIE) and density-dependent (DDE) emigration on the extinction risk of local populations and a two-patch metapopulation. (1) For completely isolated patches extinction risk increases linearly with realised emigration rates in the DIE scenario. (2) For the DDE scenario extinction risk is nearly insensitive to emigration as longs as emigration probabilities remain below ≈0.2. Survival chances are up to half an order of magnitude larger than for populations with DIE. (3) For low dispersal mortality both modes of emigration increase survival of a metapopulation by ca. one order of magnitude. (4) For high dispersal mortality only DDE can improve the global survival chances of the metapopulation. (5) With DDE individuals are only removed from a population at high population density and the risk of extinction due to demographic stochasticity is thus much smaller compared to the DIE scenario.With density-dependent emigration prospects of metapopulations survival may thus be much higher compared to a system with density-independent emigration. Consequently, the knowledge about the factors driving emigration may significantly affect our conclusions concerning the conservation status of species.     

A mechanistic model of tree competition and facilitation for Mediterranean forests: Scaling from leaf physiology to stand dynamics

Mechanistic theories of plant competition developed to explain changes in community structure and dynamics along resource availability gradients have been mostly applied to temperate forests and grasslands where light and nutrients are the two main limiting resources. In contrast, the mechanisms underlying the structure and dynamics of water-limited plant communities have been little explored. Also previous mechanistic models rely either on complex simulators, which are difficult to interpret or on simple conceptual models, which ignore too many critical details. In this study, we develop a model of stand dynamics for light and water-limited forests of intermediate complexity and we provide an analytical framework for its analyses. The model is an individual-based simulator that describes the feedback between transpiration, stomatal function and soil water dynamics with asymmetrical competition for light and water. Trees allocate carbon to three main compartments: shoot, stem and roots. We use the model to explore general patterns that may emerge across levels of biological organization from the leaf to the stand. Model predictions are consistent with a number of features of Mediterranean forests structure and dynamics. At the plant-level the leaf-based tradeoff between carbon gain and water loss expresses as a tradeoff between mortality and growth. This tradeoff explains plant morphological changes in above-ground biomass and root to shoot allocation along a water availability gradient. At the community-level, tradeoffs among carbon acquisition and water loss govern the sign of plant interactions along the gradient. Coexistence among morphological types was not observed for the range of parameters and environmental conditions explored. Overall the model provides an unifying explanation for the observed changes in the sign of plant to plant interactions along environmental gradients as well as a process-based formulation that can be linked to empirical studies.     

"The upper limits of vegetation on Mauna Loa, Hawaii": a 50th-anniversary reassessment

In January 1958, a survey of alpine flora was conducted along a recently constructed access road across the upper volcanic slopes of Mauna Loa, Hawaii (2525-3397 m). Only five native Hawaiian species were encountered on sparsely vegetated historic and prehistoric lava flows adjacent to the roadway. A resurvey of roadside flora in 2008 yielded a more than fourfold increase to 22 species, including nine native species not previously recorded. Eight new alien species have now invaded this alpine environment, although exclusively limited to a few individuals in ruderal habitat along the roadway. Alternative explanations for species invasion and altitudinal change over the past 50 years are evaluated: (1) changes related to continuing primary succession on ameliorating (weathering) young lava substrates; (2) local climate change; and (3) road improvements and increased vehicular access which promote enhanced car-borne dispersal of alien species derived from the expanding pool of potential colonizers naturalized on the island in recent decades. Unlike alpine environments in temperate latitudes, the energy component (warming) in climate change on Mauna Loa does not appear to be the unequivocal driver of plant invasion and range extension. Warming may be offset by other climate change factors including rainfall and evapotranspiration.     

From population sources to sieves: the matrix alters host-parasitoid source-sink structure

Field experiments that examine the impact of immigration, emigration, or landscape structure (e.g., the composition of the matrix) on the source sink dynamics of fragmented populations are scarce. Here, planthoppers (Prokelisia crocea) and egg parasitoids (Anagrus columbi) were released among host-plant patches that varied in structural (caged, isolated, or in a network of other patches) and functional (mudflat matrix that impedes dispersal vs. brome-grass matrix that facilitates dispersal) connectivity. Planthoppers and parasitoids on caged patches exhibited density-dependent growth rates, achieved high equilibrium densities, and rarely went extinct. Therefore, experimental cordgrass patches were classified as population sources. Because access to immigrants did not result in elevated population densities, source populations were not also pseudosinks, i.e., patches whose densities occur above carrying capacity due to high immigration. Planthoppers and parasitoids in open patches in mudflat had dynamics similar to those in caged patches, but went extinct in 4-5 generations in open patches in brome. Brome-embedded patches leaked emigrants at a rate that exceeded the gains from reproduction and immigration; populations of this sort are known as population sieves. For species whose suitable patches are becoming smaller and more isolated as a result of increased habitat fragmentation, emigration losses are likely to become paramount, a condition favoring the formation of population sieves. An increase in the proportion of patches that are sieves is predicted to destabilize regional population dynamics.     

Role of Light Availability and Dispersal in Exotic Plant Invasion along Roads and Streams in the H. J. Andrews Experimental Forest, Oregon

Abstract: We examined the roles of dispersal mechanism, a biological barrier; light availability, an environmental barrier; and level of disturbance, a physical barrier, in explaining the spatial patterns of exotic plant species along road and stream segments in a forest landscape in the western Cascade Range of Oregon (U.S.A). The presence or absence of 21 selected exotic plant species and light levels were observed along 0.3- to 1.0-km transects within four habitat types. Each habitat represented a different level of disturbance: high-use roads, low-use roads, abandoned roads, and streams in the H. J. Andrews Experimental Forest. Nearly 300 50 × 2–m sampling units were surveyed along five transects in each habitat type. We used ordination (nonmetric multidimensional scaling) and logistic regression to analyze data. All of the nearly 200 sampling units along roads with high and low levels of vehicle traffic contained at least one exotic plant species, and some contained as many as 14. Streams that were most recently disturbed by floods 20–30 years ago and abandoned spur roads with no traffic for 20–40 years also had numerous exotic species. Roads and streams apparently serve multiple functions that enhance exotic species invasion in this landscape: they act as corridors or agents for dispersal, provide suitable habitat, and contain reservoirs of propagules for future episodes of invasion. Species-specific dispersal mechanisms, habitat characteristics, and disturbance history each explain some, but not all, of the patterns of exotic species invasion observed in this study.     

Gradient forests: calculating importance gradients on physical predictors

In ecological analyses of species and community distributions there is interest in the nature of their responses to environmental gradients and in identifying the most important environmental variables, which may be used for predicting patterns of biodiversity. Methods such as random forests already exist to assess predictor importance for individual species and to indicate where along gradients abundance changes. However, there is a need to extend these methods to whole assemblages, to establish where along the range of these gradients the important compositional changes occur, and to identify any important thresholds or change points. We develop such a method, called "gradient forest," which is an extension of the random forest approach. By synthesizing the cross-validated R2 and accuracy importance measures from univariate random forest analyses across multiple species, sampling devices, and surveys, gradient forest obtains a monotonic function of each predictor that represents the compositional turnover along the gradient of the predictor. When applied to a synthetic data set, the method correctly identified the important predictors and delineated where the compositional change points occurred along these gradients. Application of gradient forest to a real data set from part of the Great Barrier Reef identified mud fraction of the sediment as the most important predictor, with highest compositional turnover occurring at mud fraction values around 25%, and provided similar information for other predictors. Such refined information allows for more accurate capturing of biodiversity patterns for the purposes of bioregionalization, delineation of protected areas, or designing of biodiversity surveys.     

Identifying robust strategies for assisted migration in a competitive stochastic metacommunity

Assisted migration (AM) is the translocation of species beyond their historical range to locations that are expected to be more suitable under future climate change. However, a relocated population may fail to establish in its donor community if there is high uncertainty in decision-making, climate, and interactions with the recipient ecological community. To quantify the benefit to persistence and risk of establishment failure of AM under different management scenarios (e.g., choosing target species, proportion of population to relocate, and optimal location to relocate), we built a stochastic metacommunity model to simulate several species reproducing, dispersing, and competing on a temperature gradient as temperature increases over time. Without AM, the species were vulnerable to climate change when they had low population sizes, short dispersal, and strong poleward competition. When relocating species that exemplified these traits, AM increased the long-term persistence of the species most when relocating a fraction of the donor population, even if the remaining population was very small or rapidly declining. This suggests that leaving behind a fraction of the population could be a robust approach, allowing managers to repeat AM in case they move the species to the wrong place and at the wrong time, especially when it is difficult to identify a species’ optimal climate. We found that AM most benefitted species with low dispersal ability and least benefited species with narrow thermal tolerances, for which AM increased extinction risk on average. Although relocation did not affect the persistence of nontarget species in our simple competitive model, researchers will need to consider a more complete set of community interactions to comprehensively understand invasion potential.     

Using demography and movement behavior to predict range expansion of the southern sea otter

In addition to forecasting population growth, basic demographic data combined with movement data provide a means for predicting rates of range expansion. Quantitative models of range expansion have rarely been applied to large vertebrates, although such tools could be useful for restoration and management of many threatened but recovering populations. Using the southern sea otter (Enhydra lutris nereis) as a case study, we utilized integro-difference equations in combination with a stage-structured projection matrix that incorporated spatial variation in dispersal and demography to make forecasts of population recovery and range recolonization. In addition to these basic predictions, we emphasize how to make these modeling predictions useful in a management context through the inclusion of parameter uncertainty and sensitivity analysis. Our models resulted in hind-cast (1989-2003) predictions of net population growth and range expansion that closely matched observed patterns. We next made projections of future range expansion and population growth, incorporating uncertainty in all model parameters, and explored the sensitivity of model predictions to variation in spatially explicit survival and dispersal rates. The predicted rate of southward range expansion (median = 5.2 km/yr) was sensitive to both dispersal and survival rates; elasticity analysis indicated that changes in adult survival would have the greatest potential effect on the rate of range expansion, while perturbation analysis showed that variation in subadult dispersal contributed most to variance in model predictions. Variation in survival and dispersal of females at the south end of the range contributed most of the variance in predicted southward range expansion. Our approach provides guidance for the acquisition of further data and a means of forecasting the consequence of specific management actions. Similar methods could aid in the management of other recovering populations.     

元江流域干热河谷草地植物群落结构特征与相似性分析

选择元江干热河谷典型草地群落,采用标准样地调查法,对流域上、中、下游干热河谷草地植物群落的分布特征、群落结构、种群习性等特征开展研究,结果表明：（1）元江流域上游祥云和中下游红河干热河谷草地群落中出现的植物种类最多,元江流域干热河谷草地植物群落是以灌木为主、多年生草本为辅的结构类型。（2）元江流域干热河谷草地群落中,扭黄茅（Heteropogon contortus）为优势种群,次优种群随流域环境的改变而不同。（3）元江流域干热河谷草地植物群落之间的相似性系数均在0.00～0.25之间,处于极不相似水平,种群的扩散受流域河谷通道作用影响较小。（4）种群的扩散和定居受海拔高度的影响,随着海拔高度的变化,植物群落的组成也发生变化,形成不同的群落结构。     

Modeling the beta diversity of coral reefs

Quantifying the beta diversity (species replacement along spatiotemporal gradients) of ecosystems is important for understanding and conserving patterns of biodiversity. However, virtually all studies of beta diversity focus on one-dimensional transects orientated along a specific environmental gradient that is defined a priori. By ignoring a second spatial dimension and the associated changes in species composition and environmental gradients, this approach may provide limited insight into the full pattern of beta diversity. Here, we use remotely sensed imagery to quantify beta diversity continuously, in two dimensions, and at multiple scales across an entire tropical marine seascape. We then show that beta diversity can be modeled (0.852 > or = r2 > or = 0.590) at spatial scales between 0.5 and 5.0 km2, using the environmental variables of mean and variance of depth and wave exposure. Beta diversity, quantified within a "window" of a given size, is positively correlated to the range of environmental conditions within that window. For example, beta diversity increases with increasing variance of depth. By analyzing such relationships across seascapes, this study provides a framework for a range of disparate coral reef literature including studies of zonation, diversity, and disturbance. Using supporting evidence from soft-bottom communities, we hypothesize that depth will be an important variable for modeling beta diversity in a range of marine systems. We discuss the implications of our results for the design of marine reserves.     

Urbanized landscapes favored by fig-eating birds increase invasive but not native juvenile strangler fig abundance

Propagule pressure can determine the success or failure of invasive plant range expansion. Range expansion takes place at large spatial scales, often encompassing many types of land cover, yet the effect of landscape context on propagule pressure remains largely unknown. Many studies have reported a positive correlation between invasive plant abundance and human land use; increased propagule pressure in these landscapes may be responsible for this correlation. We tested the hypothesis that increased rates of seed dispersal by fig-eating birds, which are more common in urban habitats, result in an increase in invasive strangler fig abundance in landscapes dominated by human land use. We quantified abundance of an invasive species (Ficus microcarpa) and a native species (F. aurea) of strangler fig in plots spanning the entire range of human land use in South Florida, USA, from urban parking lots to native forest. We then compared models that predicted juvenile fig abundance based on distance to adult fig seed sources and fig-eating bird habitat quality with models that lacked one or both of these terms. The best model for juvenile invasive fig abundance included both distance to adult and fig-eating bird habitat terms, suggesting that landscape effects on invasive fig abundance are mediated by seed-dispersing birds. In contrast, the best model for juvenile native fig abundance included only presence/absence of adults, suggesting that distance from individual adult trees may have less effect on seed limitation for a native species compared to an invasive species undergoing range expansion. However, models for both species included significant effects of adult seed sources, implying that juvenile abundance is limited by seed arrival. This result was corroborated by a seed addition experiment that indicated that both native and invasive strangler figs were strongly seed limited. Understanding how landscape context affects the mechanisms of plant invasion may lead to better management techniques. Our results suggest that prioritizing removal of adult trees in sites with high fig-eating bird habitat may be the most effective method to control F. microcarpa abundance.     

Blue Tits use fledgling quantity and quality as public information in breeding site choice

Public information (PI), which is the information that can be derived from the behavior and performance of conspecifics, has been demonstrated to be used in many fitness-enhancing decisions. In the context of breeding habitat choice, PI use has been called "habitat copying." We experimentally tested the existence of habitat copying in the Blue Tit (Cyanistes caeruleus), a nonmigratory, short-lived hole-nesting bird. We manipulated the mean number of fledglings raised locally (quantity) and their condition (quality) as components of PI by transferring nestlings from Decreased (D) patches to Increased (I) patches. Our manipulations caused a negative relationship between fledgling quantity and quality that does not exist naturally: I patches had a higher number of fledglings that were in poorer condition, whereas D patches had a lower number in better condition. Control (C) patches, whether manipulated or not, had intermediate levels in terms of fledgling quantity and quality. Adult emigration the following year was higher from D than from C or I patches. Similarly, adult dispersal distance decreased for individuals coming from D to C to I patches. This suggests that resident breeders rely mainly on fledgling quantity to make emigration decisions. Emigration patterns of juveniles did not vary in relation to our patch manipulation. Immigration rates were higher and similar in I and D patches than in C patches. Hence, immigrant Blue Tits seem to rely on one of the manipulated components of PI and are insensitive to the discrepancy between fledgling quantity and quality. This shows that even nonmigratory species, such as Blue Tits, may use PI in their dispersal decisions but weigh its components differently for emigration and immigration. Differences among species in the importance of PI in breeding habitat choices may be explained by differences in life histories.     

Herbivory limits recruitment in an old-field seed addition experiment

Environmental variability can promote coexistence by creating establishment sites for rare plants, but low diversity in anthropogenic grasslands suggests that this variability may be eliminated (homogenization hypothesis) or inaccessible (barrier hypothesis). We explore these alternatives on the northern Great Plains, where 11 million hectares have been transformed by multiple environmental changes, but the causes of species loss are unclear. In a degraded grassland, we increased environmental variability by manipulating competition and herbivory along gradients of fertility and disturbance, and we circumvented dispersal barriers by adding 1.2 million seeds of five functionally distinct species at varying densities. The experiment ended after 12 weeks due to the direct and indirect effects of unapparent small native herbivores, which were barriers to population establishment by the added species. The direct cause of recruitment failure was browsing. The indirect cause was associated with competition from invasive plants that appeared to be more tolerant or resistant to herbivory. Variability in fertility, disturbance, propagule pressure, and competition had relatively minor impacts on colonization by the added species because herbivores controlled recruitment in most environments. Recruitment outside the herbivore exclosures was mostly by unpalatable exotics, suggesting a possible link between invasion success and herbivore resistance for some introduced plants.