Racing against change: understanding dispersal and persistence to improve species' conservation prospects

Climate change is contributing to the widespread redistribution, and increasingly the loss, of species. Geographical range shifts among many species were detected rapidly after predictions of the potential importance of climate change were specified 35 years ago: species are shifting their ranges towards the poles and often to higher elevations in mountainous areas. Early tests of these predictions were largely qualitative, though extraordinarily rapid and broadly based, and statistical tests distinguishing between climate change and other global change drivers provided quantitative evidence that climate change had already begun to cause species’ geographical ranges to shift. I review two mechanisms enabling this process, namely development of approaches for accounting for dispersal that contributes to range expansion, and identification of factors that alter persistence and lead to range loss. Dispersal in the context of range expansion depends on an array of processes, like population growth rates in novel environments, rates of individual species movements to new locations, and how quickly areas of climatically tolerable habitat shift. These factors can be tied together in well-understood mathematical frameworks or modelled statistically, leading to better prediction of extinction risk as climate changes. Yet, species'' increasing exposures to novel climate conditions can exceed their tolerances and raise the likelihood of local extinction and consequent range losses. Such losses are the consequence of processes acting on individuals, driven by factors, such as the growing frequency and severity of extreme weather, that contribute local extinction risks for populations and species. Many mechanisms can govern how species respond to climate change, and rapid progress in global change research creates many opportunities to inform policy and improve conservation outcomes in the early stages of the sixth mass extinction.     

Seed dispersal of fleshy-fruited invasive plants by birds: contributing factors and management options

The ecology of seed dispersal by vertebrates has been investigated extensively over recent decades, yet only limited research has been conducted on how suites of invasive plants and frugivorous birds interact. In this review, we examine how plant fruit traits (morphology, colour and display, nutritional quality, accessibility and phenology), avian traits (fruit handling techniques, gut passage time and effect, bird movements and social behaviour and dietary composition) and landscape structure (fruit neighbourhood, habitat loss and fragmentation and perch tree effects) affect frugivory and seed dispersal in invasive plants. This functional approach could be used to develop generic models of seed dispersal distributions for suites of invasive plant species and improve management efficiencies. Four broad research approaches are described that could direct management of bird‐dispersed invasive plants at the landscape scale, by manipulating dispersal. First, research is needed to quantify the effect of biological control agents on dispersal, particularly how changes in fruit production and/or quality affect fruit choice by frugivores, dispersal distributions of seed and post‐dispersal processes. Second, we explore how seed dispersal could be directed, such as by manipulating perch structures and/or vegetation density to attract frugivorous birds after they have been foraging on invasive plant fruits. Third, the major sources of seed spread could be identified and removed (i.e. targeting core or satellite infestations, particular habitats and creating barrier zones). Fourth, alternative food resources could be provided for frugivores, to replace fruits of invasive plants, and their use quantified.     

Dispersal evolution in temporally variable environments: implications for plant range dynamics

Dispersal—the movement of an individual from the site of birth to a different site for reproduction—is an ecological and evolutionary driver of species ranges that shapes patterns of colonization, connectivity, gene flow, and adaptation. In plants, the traits that influence dispersal often vary within and among species, are heritable, and evolve in response to the fitness consequences of moving through heterogeneous landscapes. Spatial and temporal variation in the quality and quantity of habitat are important sources of selection on dispersal strategies across species ranges. While recent reviews have evaluated the interactions between spatial variation in habitat and dispersal dynamics, the extent to which geographic variation in temporal variability can also shape range-wide patterns in dispersal traits has not been synthesized. In this paper, we summarize key predictions from metapopulation models that evaluate how dispersal evolves in response to spatial and temporal habitat variability. Next, we compile empirical data that quantify temporal variability in plant demography and patterns of dispersal trait variation across species ranges to evaluate the hypothesis that higher temporal variability favors increased dispersal at plant range limits. We found some suggestive evidence supporting this hypothesis while more generally identifying a major gap in empirical work evaluating plant metapopulation dynamics across species ranges and geographic variation in dispersal traits. To address this gap, we propose several future research directions that would advance our understanding of the interplay between spatiotemporal variability and dispersal trait variation in shaping the dynamics of current and future species ranges.     

The importance of long-distance dispersal in biodiversity conservation

Dispersal is universally considered important for biodiversity conservation. However, the significance of long‐ as opposed to short‐distance dispersal is insufficiently recognized in the conservation context. Long‐distance dispersal (LDD) events, although typically rare, are crucial to population spread and to maintenance of genetic connectivity. The main threats to global biodiversity involve excessive LDD of elements alien to ecosystems and insufficient dispersal of native species, for example, because of habitat fragmentation. In this paper, we attempt to bridge the gap in the treatment of LDD by reviewing the conservation issues for which LDD is most important. We then demonstrate how taking LDD into consideration can improve conservation management decisions.     

Range expansion by invasive marine algae: rates and patterns of spread at a regional scale

Aim  Introduced macroalgae are widespread in the world's oceans and, despite increasing awareness and attempts to limit the phenomenon, the number of species introductions in coastal waters has increased exponentially over time. Little is known about the rates and mechanisms of spread, even among species that have received the most attention. We compare patterns of range expansion for nine species of invasive algae across eight geographic regions.
Location  World-wide.
Methods  We compiled records of introduced algae from the scientific literature, herbaria, and by contacting experts to reconstruct chronologies for 22 algal invasions. These were used to map patterns of spread at a regional scale (thousands of km).
Results  Range size tended to increase linearly with time, often after an initial lag. Range expansion occurred at rates of tens to hundreds of kilometres per year, often with large infrequent increases. Rates of range expansion differed significantly between species within the same region, and between regions for the same species.
Main conclusions  Our results suggest that anthropogenic vectors likely play a key role in the spread of introduced macroalgae at a regional scale, although natural long-distance dispersal also may be important for some species. The lack of consistency in the rates within individual species and regions suggests that multiple interacting factors (e.g. algal traits, characteristics of invaded communities, environmental conditions and anthropogenic activities) determine where propagules of introduced algae are delivered and whether they become established.     

Mutation surfing and the evolution of dispersal during range expansions

A growing body of empirical evidence demonstrates that at an expanding front, there can be strong selection for greater dispersal propensity, whereas recent theory indicates that mutations occurring towards the front of a spatially expanding population can sometimes ‘surf’ to high frequency and spatial extent. Here, we consider the potential interplay between these two processes: what role may mutation surfing play in determining the course of dispersal evolution and how might dispersal evolution itself influence mutation surfing? Using an individual‐based coupled‐map lattice model, we first run simulations to determine the fate of dispersal mutants that occur at an expanding front. Our results highlight that mutants that have a slightly higher dispersal propensity than the wild type always have a higher survival probability than those mutants with a dispersal propensity lower than, or very similar to, the wild type. However, it is not always the case that mutants with very high dispersal propensity have the greatest survival probability. When dispersal mortality is high, mutants of intermediate dispersal survive most often. Interestingly, the rate of dispersal that ultimately evolves at an expanding front is often substantially higher than that which confers a novel mutant with the greatest probability of survival. Second, we run a model in which we allow dispersal to evolve over the course of a range expansion and ask how the fate of a neutral or nonneutral mutant depends upon when and where during the expansion it arises. These simulations highlight that the success of a neutral mutant depends upon the dispersal genotypes that it is associated with. An important consequence of this is that novel mutants that arise at the front of an expansion, and survive, typically end up being associated with more dispersive genotypes than the wild type. These results offer some new insights into causes and the consequences of dispersal evolution during range expansions, and the methodology we have employed can be readily extended to explore the evolutionary dynamics of other life history characteristics.     

Recruitment trade-offs and the evolution of dispersal mechanisms in plants

In this study we place seed size vs. seed number trade-offs in the context of plant dispersal ability. The objective was to suggest explanations for the evolution of different seed dispersal mechanisms, in particular fleshy fruits, wind dispersal and the maintenance of unassisted dispersal. We suggest that selection for improved dispersal may act either by increasing the intercept of a dispersal curve (log seed number vs. distance) or by flattening the slope of the curve. 'Improved dispersal' is defined as a marginal increase in the number of recruits sited at some (arbitrary) distance away from the parent plant. Increasing the intercept of the dispersal curve, i.e. producing more seeds, is associated with a reduction in seed size, which in turn affects the recruitment ability, provided that this ability is related to seed size. If recruitment is related to seed size there will be a recruitment cost of evolving increased seed production. On the other hand, a flattening of the slope by evolving dispersal attributes is likely to be associated with a fecundity cost. An exception is wind dispersal where smaller (and hence more numerous) seeds may lead to more efficient dispersal. We derive two main predictions: If recruitment is strongly related to seed size, selection for improved dispersal acts on the slope of the dispersal curve, i.e. by favouring evolution of dispersal attributes on seeds or fruits. If, on the other hand, recruitment is only weakly related to seed size (or not related, or negatively related), selection for improved dispersal favours increased seed production. Despite its simplicity, the model suggests explanations for (i) why so many plant species lack special seed dispersal attributes, (ii) differences in dispersal spectra among plant communities, and (iii) adaptive radiation in seed size and dispersal attributes during angiosperm evolution. This revised version was published online in July 2006 with corrections to the Cover Date.     

Species richness of herbivores on exotic host plants increases with time since introduction of the host

Aim Species richness of insect herbivores feeding on exotic plants increases with abundance as well as range size of the host in the area of introduction. The formation of these herbivore assemblages requires a certain amount of time, and the richness of insect faunas should also increase with the length of time an exotic plant has been present in the introduced range. Location Central Europe. Methods We analysed the variation in species richness of leaf‐chewing Lepidoptera larvae and sap‐sucking Auchenorrhyncha (Hemiptera) associated with 103 exotic woody plant species in Germany in relation to time since introduction, range size, growth form (trees versus shrubs), biogeographical origin (distance from Central Europe) and taxonomic isolation of the host plant (presence or absence of a native congener in the introduced area). Results Using simple correlation analyses we found for Lepidoptera and Auchenorrhyncha that species richness increased with time since introduction of the host plant. For the Lepidoptera the increase of species richness with time since introduction remained significant even after removing the effects of all other independent variables. Main conclusions Our results provide some evidence that assemblages of insects on exotic plants do not reach saturation within a time scale of few hundred years. This contrasts with previous findings for crop plants.     

Long range dispersal and spatial pattern formation in biological invasions

In this paper we explore the consequences of long distance dispersal in biological invasion processes through simulations using a recently developed cellular automaton model. We show that long distance dispersal generate characteristic spatial patterns with several stationary scale-invariant properties. In particular, the patterns display a main patch around the focus of spread, with a fractal border structure whose fractal dimension contains information about the main statistical properties of the dispersal mechanism. Our results are in agreement with field data of spread of invaders with long distance dispersal mechanisms.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

Native and naturalized range size in Pinus: relative importance of biogeography,introduction effort and species traits

Aim Pine trees (genus Pinus) represent an ancient lineage, naturally occurring almost exclusively in the Northern Hemisphere, but introduced and widely naturalized in both hemispheres. As large trees of interest to forestry, they attract much attention and their distribution is well documented in both indigenous and naturalized ranges. This creates an opportunity to analyse the relationship between indigenous and naturalized range sizes in the context of different levels of human usage, biological traits and the characteristics of the environments of origin. Location Global. Methods We combined and expanded pre‐existing data sets for pine species distributions and pine species traits, and used a variety of regression techniques (including generalized additive models and zero‐inflated Poisson models) to assess which variables explained naturalized and indigenous range sizes. Results Indigenous and naturalized range sizes are positively correlated but there are many notable exceptions. Some species have large indigenous ranges but small or no naturalized ranges, whereas others have small indigenous ranges, but have naturalized in many regions. Indigenous range is correlated to factors such as seed size (?), age at first reproduction (?), and latitude (+, supporting Rapoport's rule), but also to the extent of coverage of species in the forestry literature (+). Naturalized range size is strongly influenced by the extent of coverage of species in the forestry literature (+), a proxy for propagule pressure. Naturalization was also influenced by average elevation in the indigenous range (?) and age at first reproduction (?). Main conclusions The macroecological and evolutionary pressures facing plant groups are not directly transferable between indigenous and naturalized ranges. In particular, there are strong biases in species naturalization and expansion in invasive ranges that are unrelated to factors determining indigenous range size. At least for Pinus, a new set of macroecological patterns are emerging which are profoundly influenced by humans.     

Fitness declines towards range limits and local adaptation to climate affect dispersal evolution during climate‐induced range shifts

Dispersal ability will largely determine whether species track their climatic niches during climate change, a process especially important for populations at contracting (low‐latitude/low‐elevation) range limits that otherwise risk extinction. We investigate whether dispersal evolution at contracting range limits is facilitated by two processes that potentially enable edge populations to experience and adjust to the effects of climate deterioration before they cause extinction: (i) climate‐induced fitness declines towards range limits and (ii) local adaptation to a shifting climate gradient. We simulate a species distributed continuously along a temperature gradient using a spatially explicit, individual‐based model. We compare range‐wide dispersal evolution during climate stability vs. directional climate change, with uniform fitness vs. fitness that declines towards range limits (RLs), and for a single climate genotype vs. multiple genotypes locally adapted to temperature. During climate stability, dispersal decreased towards RLs when fitness was uniform, but increased when fitness declined towards RLs, due to highly dispersive genotypes maintaining sink populations at RLs, increased kin selection in smaller populations, and an emergent fitness asymmetry that favoured dispersal in low‐quality habitat. However, this initial dispersal advantage at low‐fitness RLs did not facilitate climate tracking, as it was outweighed by an increased probability of extinction. Locally adapted genotypes benefited from staying close to their climate optima; this selected against dispersal under stable climates but for increased dispersal throughout shifting ranges, compared to cases without local adaptation. Dispersal increased at expanding RLs in most scenarios, but only increased at the range centre and contracting RLs given local adaptation to climate.     

MigClim: Predicting plant distribution and dispersal in a changing climate

Aim Many studies have forecasted the possible impact of climate change on plant distributions using models based on ecological niche theory, but most of them have ignored dispersal‐limitations, assuming dispersal to be either unlimited or null. Depending on the rate of climatic change, the landscape fragmentation and the dispersal capabilities of individual species, these assumptions are likely to prove inaccurate, leading to under‐ or overestimation of future species distributions and yielding large uncertainty between these two extremes. As a result, the concepts of ‘potentially suitable’ and ‘potentially colonizable’ habitat are expected to differ significantly. To quantify to what extent these two concepts can differ, we developed Mig Clim, a model simulating plant dispersal under climate change and landscape fragmentation scenarios. Mig Clim implements various parameters, such as dispersal distance, increase in reproductive potential over time, landscape fragmentation or long‐distance dispersal. Location Western Swiss Alps. Methods Using our Mig Clim model, several simulations were run for two virtual species by varying dispersal distance and other parameters. Each simulation covered the 100‐year period 2001–2100 and three different IPCC‐based temperature warming scenarios were considered. Results of dispersal‐limited projections were compared with unlimited and no‐dispersal projections. Results Our simulations indicate that: (1) using realistic parameter values, the future potential distributions generated using Mig Clim can differ significantly (up to more than 95% difference in colonized surface) from those that ignore dispersal; (2) this divergence increases under more extreme climate warming scenarios and over longer time periods; and (3) the uncertainty associated with the warming scenario can be as large as the one related to dispersal parameters. Main conclusions Accounting for dispersal, even roughly, can importantly reduce uncertainty in projections of species distribution under climate change scenarios.     

Determinants of fine‐scale homogenization and differentiation of native freshwater fish faunas in a Mediterranean Basin: implications for conservation

Aim Increasing threats to freshwater biodiversity are rapidly changing the distinctiveness of regional species pools and local assemblages. Biotic homogenization/differentiation processes are threatening the integrity and persistence of native biodiversity patterns at a range of spatial scales and pose a challenge for effective conservation planning. Here, we evaluate the extent and determinants of fine‐scale alteration in native freshwater fish assemblages among stream reaches throughout a large river basin and consider the implications of these changes for the long‐term conservation of native fishes. Location Guadiana River basin (South‐Western Iberian Peninsula). Methods We quantified the magnitude of change in compositional similarity between observed and reference assemblages and its potential effect on natural patterns of compositional distinctiveness. Reference assemblages were defined as the native species expected to occur naturally (in absence of anthropogenic alterations) and were reconstructed using a multivariate adaptive regression splines predictive model. We also evaluated the role of habitat degradation and introduced species as determinants of biotic homogenization/differentiation. Results We found a significant trend towards homogenization for native fish assemblages. Changes in native fish distributions led to the loss of distinctiveness patterns along natural environmental gradients. Introduced species were the most important factor explaining the homogenization process. Homogenization of native assemblages was stronger in areas close to reservoirs and in lowland reaches where introduced species were more abundant. Main conclusions The implementation of efficient conservation for the maintenance of native fish diversity is seriously threatened by the homogenization processes. The identification of priority areas for conservation is hindered by the fact that the most diverse communities are vanishing, which would require the selection of broader areas to adequately protect all the species. Given the principal role that introduced species play in the homogenization process and their relation with reservoirs, special attention must be paid to mitigating or preventing these threats.