Ecological niche models of invasive seaweeds

Ecological niche models (ENMs) are commonly used to calculate habitat suitability from species’ occurrence and macroecological data. In invasive species biology, ENMs can be applied to anticipate whether invasive species are likely to establish in an area, to identify critical routes and arrival points, to build risk maps and to predict the extent of potential spread following an introduction. Most studies using ENMs focus on terrestrial organisms and applications in the marine realm are still relatively rare. Here, we review some common methods to build ENMs and their application in seaweed invasion biology. We summarize methods and concepts involved in the development of niche models, show examples of how they have been applied in studies on algae and discuss the application of ENMs in invasive algae research and to predict effects of climate change on seaweed distributions.     

Meta‐analyzing the likely cross‐species responses to climate change

Ecological Niche Models (ENMs) have different performances in predicting potential geographic distributions. Here we meta‐analyzed the likely effects of climate change on the potential geographic distribution of 1,205 bird species from the Neotropical region, modeled using eight ENMs and three Atmosphere‐Ocean General Circulation Models (AOGCM). We considered the variability in ENMs performance to estimate a weighted mean difference between potential geographic distributions for baseline and future climates. On average, potential future ranges were projected to be from 25.7% to 44.5% smaller than current potential ranges across species. However, we found that 0.2% to 18.3% of the total variance in range shifts occurred “within species” (i.e., owing to the use of different modeling techniques and climate models) and 81.7% to 99.8% remained between species (i.e., it could be explained by ecological correlates). Using meta‐analytical techniques akin to regression, we also showed that potential range shifts are barely predicted by bird biological traits. We demonstrated that one can combine and reduce species‐specific effects with high uncertainty in ENMs and also explore potential causes of climate change effect on species using meta‐analytical tools. We also highlight that the search for powerful correlates of climate change‐induced range shifts can be a promising line of investigation.     

Eco‐evolution on the edge during climate change

We urgently need to predict species responses to climate change to minimize future biodiversity loss and ensure we do not waste limited resources on ineffective conservation strategies. Currently, most predictions of species responses to climate change ignore the potential for evolution. However, evolution can alter species ecological responses, and different aspects of evolution and ecology can interact to produce complex eco‐evolutionary dynamics under climate change. Here we review how evolution could alter ecological responses to climate change on species warm and cool range margins, where evolution could be especially important. We discuss different aspects of evolution in isolation, and then synthesize results to consider how multiple evolutionary processes might interact and affect conservation strategies. On species cool range margins, the evolution of dispersal could increase range expansion rates and allow species to adapt to novel conditions in their new range. However, low genetic variation and genetic drift in small range‐front populations could also slow or halt range expansions. Together, these eco‐evolutionary effects could cause a three‐step, stop‐and‐go expansion pattern for many species. On warm range margins, isolation among populations could maintain high genetic variation that facilitates evolution to novel climates and allows species to persist longer than expected without evolution. This ‘evolutionary extinction debt’ could then prevent other species from shifting their ranges. However, as climate change increases isolation among populations, increasing dispersal mortality could select for decreased dispersal and cause rapid range contractions. Some of these eco‐evolutionary dynamics could explain why many species are not responding to climate change as predicted. We conclude by suggesting that resurveying historical studies that measured trait frequencies, the strength of selection, or heritabilities could be an efficient way to increase our eco‐evolutionary knowledge in climate change biology.     

Different climatic envelopes among invasive populations may lead to underestimations of current and future biological invasions

Aim We explore the impact of calibrating ecological niche models (ENMs) using (1) native range (NR) data versus (2) entire range (ER) data (native and invasive) on projections of current and future distributions of three Hieracium species. Location H. aurantiacum, H. murorum and H. pilosella are native to Europe and invasive in Australia, New Zealand and North America. Methods Differences among the native and invasive realized climatic niches of each species were quantified. Eight ENMs in BIOMOD were calibrated with (1) NR and (2) ER data. Current European, North American and Australian distributions were projected. Future Australian distributions were modelled using four climate change scenarios for 2030. Results The invasive climatic niche of H. murorum is primarily a subset of that expressed in its native range. Invasive populations of H. aurantiacum and H. pilosella occupy different climatic niches to those realized in their native ranges. Furthermore, geographically separate invasive populations of these two species have distinct climatic niches. ENMs calibrated on the realized niche of native regions projected smaller distributions than models incorporating data from species’ entire ranges, and failed to correctly predict many known invasive populations. Under future climate scenarios, projected distributions decreased by similar percentages, regardless of the data used to calibrate ENMs; however, the overall sizes of projected distributions varied substantially. Main conclusions This study provides quantitative evidence that invasive populations of Hieracium species can occur in areas with different climatic conditions than experienced in their native ranges. For these, and similar species, calibration of ENMs based on NR data only will misrepresent their potential invasive distribution. These errors will propagate when estimating climate change impacts. Thus, incorporating data from species’ entire distributions may result in a more thorough assessment of current and future ranges, and provides a closer approximation of the elusive fundamental niche.     

Both life‐history plasticity and local adaptation will shape range‐wide responses to climate warming in the tundra plant Silene acaulis

Many predictions of how climate change will impact biodiversity have focused on range shifts using species‐wide climate tolerances, an approach that ignores the demographic mechanisms that enable species to attain broad geographic distributions. But these mechanisms matter, as responses to climate change could fundamentally differ depending on the contributions of life‐history plasticity vs. local adaptation to species‐wide climate tolerances. In particular, if local adaptation to climate is strong, populations across a species’ range—not only those at the trailing range edge—could decline sharply with global climate change. Indeed, faster rates of climate change in many high latitude regions could combine with local adaptation to generate sharper declines well away from trailing edges. Combining 15 years of demographic data from field populations across North America with growth chamber warming experiments, we show that growth and survival in a widespread tundra plant show compensatory responses to warming throughout the species’ latitudinal range, buffering overall performance across a range of temperatures. However, populations also differ in their temperature responses, consistent with adaptation to local climate, especially growing season temperature. In particular, warming begins to negatively impact plant growth at cooler temperatures for plants from colder, northern populations than for those from warmer, southern populations, both in the field and in growth chambers. Furthermore, the individuals and maternal families with the fastest growth also have the lowest water use efficiency at all temperatures, suggesting that a trade‐off between growth and water use efficiency could further constrain responses to forecasted warming and drying. Taken together, these results suggest that populations throughout species’ ranges could be at risk of decline with continued climate change, and that the focus on trailing edge populations risks overlooking the largest potential impacts of climate change on species’ abundance and distribution.     

Ecological niche models in phylogeographic studies: applications,advances and precautions

The increased availability of spatial data and methodological developments in species distribution modelling has lead to concurrent advances in phylogeography, broadening the scope of questions studied, as well as providing unprecedented insights. Given the species‐specific nature of the information provided by ecological niche models (ENMs), whether it is on the environmental tolerances of species or their estimated distribution, today or in the past, it is perhaps not surprising that ENMs have rapidly become a common tool in phylogeographic analysis. Such information is essential to phylogeographic tests that provide important biological insights. Here, we provide an overview of the different applications of ENMs in phylogeographic studies, detailing specific studies and highlighting general limitations and challenges with each application. Given that the full potential of integrating ENMs into phylogeographic cannot be realized unless the ENMs themselves are carefully applied, we provide a summary of best practices with using ENMs. Lastly, we describe some recent advances in how quantitative information from ENMs can be integrated into genetic analyses, illustrating their potential use (and key concerns with such implementations), as well as promising areas for future development.     

Plant adaptation to climate change—Where are we?

Climate change poses critical challenges for population persistence in natural communities, for agriculture and environmental sustainability, and for food security. In this review, we discuss recent progress in climatic adaptation in plants. We evaluate whether climate change exerts novel selection and disrupts local adaptation, whether gene flow can facilitate adaptive responses to climate change, and whether adaptive phenotypic plasticity could sustain populations in the short term. Furthermore, we discuss how climate change influences species interactions. Through a more in‐depth understanding of these eco‐evolutionary dynamics, we will increase our capacity to predict the adaptive potential of plants under climate change. In addition, we review studies that dissect the genetic basis of plant adaptation to climate change. Finally, we highlight key research gaps, ranging from validating gene function to elucidating molecular mechanisms, expanding research systems from model species to other natural species, testing the fitness consequences of alleles in natural environments, and designing multifactorial studies that more closely reflect the complex and interactive effects of multiple climate change factors. By leveraging interdisciplinary tools (e.g., cutting‐edge omics toolkits, novel ecological strategies, newly developed genome editing technology), researchers can more accurately predict the probability that species can persist through this rapid and intense period of environmental change, as well as cultivate crops to withstand climate change, and conserve biodiversity in natural systems.     

Predicting phenology by integrating ecology,evolution and climate science

Forecasting how species and ecosystems will respond to climate change has been a major aim of ecology in recent years. Much of this research has focused on phenology – the timing of life‐history events. Phenology has well‐demonstrated links to climate, from genetic to landscape scales; yet our ability to explain and predict variation in phenology across species, habitats and time remains poor. Here, we outline how merging approaches from ecology, climate science and evolutionary biology can advance research on phenological responses to climate variability. Using insight into seasonal and interannual climate variability combined with niche theory and community phylogenetics, we develop a predictive approach for species’ reponses to changing climate. Our approach predicts that species occupying higher latitudes or the early growing season should be most sensitive to climate and have the most phylogenetically conserved phenologies. We further predict that temperate species will respond to climate change by shifting in time, while tropical species will respond by shifting space, or by evolving. Although we focus here on plant phenology, our approach is broadly applicable to ecological research of plant responses to climate variability.     

Losing your edge: climate change and the conservation value of range‐edge populations

Populations occurring at species' range edges can be locally adapted to unique environmental conditions. From a species' perspective, range‐edge environments generally have higher severity and frequency of extreme climatic events relative to the range core. Under future climates, extreme climatic events are predicted to become increasingly important in defining species' distributions. Therefore, range‐edge genotypes that are better adapted to extreme climates relative to core populations may be essential to species' persistence during periods of rapid climate change. We use relatively simple conceptual models to highlight the importance of locally adapted range‐edge populations (leading and trailing edges) for determining the ability of species to persist under future climates. Using trees as an example, we show how locally adapted populations at species' range edges may expand under future climate change and become more common relative to range‐core populations. We also highlight how large‐scale habitat destruction occurring in some geographic areas where many species range edge converge, such as biome boundaries and ecotones (e.g., the arc of deforestation along the rainforest‐cerrado ecotone in the southern Amazonia), can have major implications for global biodiversity. As climate changes, range‐edge populations will play key roles in helping species to maintain or expand their geographic distributions. The loss of these locally adapted range‐edge populations through anthropogenic disturbance is therefore hypothesized to reduce the ability of species to persist in the face of rapid future climate change.     

Physiology in ecological niche modeling: using zebra mussel's upper thermal tolerance to refine model predictions through Bayesian analysis

Climate change and human-mediated dispersal are increasingly influencing species’ geographic distributions. Ecological niche models (ENMs) are widely used in forecasting species’ distributions, but are weak in extrapolation to novel environments because they rely on available distributional data and do not incorporate mechanistic information, such as species’ physiological response to abiotic conditions. To improve accuracy of ENMs, we incorporated physiological knowledge through Bayesian analysis. In a case study of the zebra mussel Dreissena polymorpha, we used native and global occurrences to obtain native and global models representing narrower and broader understanding of zebra mussel’ response to temperature. We also obtained thermal limit and survival information for zebra mussel from peer-reviewed literature and used the two types of information separately and jointly to calibrate native models. We showed that, compared to global models, native models predicted lower relative probability of presence along zebra mussel's upper thermal limit, suggesting the shortcoming of native models in predicting zebra mussel's response to warm temperature. We also found that native models showed improved prediction of relative probability of presence when thermal limit was used alone, and best approximated global models when both thermal limit and survival data were used. Our result suggests that integration of physiological knowledge enhances extrapolation of ENM in novel environments. Our modeling framework can be generalized for other species or other physiological limits and may incorporate evolutionary information (e.g. evolved thermal tolerance), thus has the potential to improve predictions of species’ invasive potential and distributional response to climate change.     

Genes on the edge: A framework to detect genetic diversity imperiled by climate change

Ongoing global warming is disrupting several ecological and evolutionary processes, spanning different levels of biological organization. Species are expected to shift their ranges as a response to climate change, with relevant implications to peripheral populations at the trailing and leading edges. Several studies have analyzed the exposure of species to climate change but few have explored exposure at the intraspecific level. We introduce a framework to forecast exposure to climate change at the intraspecific level. We build on existing methods by combining correlative species distribution models, a model of species range dynamics, and a model of phylogeographic interpolation. We demonstrate the framework by applying it to 20 Iberian amphibian and reptile species. Our aims were to: (a) identify which species and intraspecific lineages will be most exposed to future climate change; (b) test if nucleotide diversity at the edges of species ranges are significantly higher or lower than on the overall range; and (c) analyze if areas of higher species gain, loss, and turnover coincide with those predicted for lineages richness and nucleotide diversity. We found that about 80% of the studied species are predicted to contract their range. Within each species, some lineages were predicted to contract their range, while others were predicted to maintain or expand it. Therefore, estimating the impacts of climate change at the species level only can underestimate losses at the intraspecific level. Some species had significant high amount of nucleotide at the trailing or leading edge, or both, but we did not find a consistent pattern across species. Spatial patterns of species richness, gain, loss, and turnover were fairly concurrent with lineages richness and nucleotide diversity. Our results support the need for increased attention to intraspecific diversity regarding monitoring and conservation strategies under climate change.     

The role of climate,habitat, and species co‐occurrence as drivers of change in small mammal distributions over the past century

Species distribution models are commonly used to predict species responses to climate change. However, their usefulness in conservation planning and policy is controversial because they are difficult to validate across time and space. Here we capitalize on small mammal surveys repeated over a century in Yosemite National Park, USA, to assess accuracy of model predictions. Historical (1900–1940) climate, vegetation, and species occurrence data were used to develop single‐ and multi‐species multivariate adaptive regression spline distribution models for three species of chipmunk. Models were projected onto the current (1980–2007) environmental surface and then tested against modern field resurveys of each species. We evaluated models both within and between time periods and found that even with the inclusion of biotic predictors, climate alone is the dominant predictor explaining the distribution of the study species within a time period. However, climate was not consistently an adequate predictor of the distributional change observed in all three species across time. For two of the three species, climate alone or climate and vegetation models showed good predictive performance across time. The stability of the distribution from the past to present observed in the third species, however, was not predicted by our modeling approach. Our results demonstrate that correlative distribution models are useful in understanding species' potential responses to environmental change, but also show how changes in species‐environment correlations through time can limit the predictive performance of models.     

Modelle für den Wandel der Natur

Ecological Modelling – Scenarios of future environmental changes Climate change affects ecosystems at different levels. Changes in species phenology, distribution and interactions are today well described phenomena documenting species responses to increasing temperatures. Environmental niche models (ENMs) have developed as powerful tools to address various questions in macroecology. Aiming at a species environmental niche, statistical modelling can be employed to predict a species' potential occurrence by projecting environmental information recorded at locality records over space and time. In climate change biology, ENMs are used to identify individual species' fates as range expansions or retractions as well as features that affect the structure of species assemblages and species interactions within and across different taxonomic groups. ENMs help to promote the persistence of species by identifying spatial patterns of species richness or endangerment to target conservation priorities. Moreover, they are an essential part of risk assessments to set up preventive measures against non‐native species most likely to adversely impact native ecosystems.     

Past climate change drives current genetic structure of an endangered freshwater mussel species

Historical‐to‐recent climate change and anthropogenic disturbance affect species distributions and genetic structure. The Rio Grande watershed of the United States and Mexico encompasses ecosystems that are intensively exploited, resulting in substantial degradation of aquatic habitats. While significant anthropogenic disturbances in the Rio Grande are recent, inhospitable conditions for freshwater organisms likely existed prior to such disturbances. A combination of anthropogenic and past climate factors may contribute to current distributions of aquatic fauna in the Rio Grande basin. We used mitochondrial DNA and 18 microsatellite loci to infer evolutionary history and genetic structure of an endangered freshwater mussel, Popenaias popeii, throughout the Rio Grande drainage. We estimated spatial connectivity and gene flow across extant populations of P. popeii and used ecological niche models (ENMs) and approximate Bayesian computation (ABC) to infer its evolutionary history during the Pleistocene. structure results recovered regional and local population clusters in the Rio Grande. ENMs predicted drastic reductions in suitable habitat during the last glacial maximum. ABC analyses suggested that regional population structure likely arose in this species during the mid‐to‐late Pleistocene and was followed by a late Pleistocene population bottleneck in New Mexico populations. The local population structure arose relatively recently, perhaps due to anthropogenic factors. Popenaias popeii, one of the few freshwater mussel species native to the Rio Grande basin, is a case study for understanding how both geological and anthropogenic factors shape current population genetic structure. Conservation strategies for this species should account for the fragmented nature of contemporary populations.     

Genetically informed ecological niche models improve climate change predictions

We examined the hypothesis that ecological niche models (ENMs) more accurately predict species distributions when they incorporate information on population genetic structure, and concomitantly, local adaptation. Local adaptation is common in species that span a range of environmental gradients (e.g., soils and climate). Moreover, common garden studies have demonstrated a covariance between neutral markers and functional traits associated with a species’ ability to adapt to environmental change. We therefore predicted that genetically distinct populations would respond differently to climate change, resulting in predicted distributions with little overlap. To test whether genetic information improves our ability to predict a species’ niche space, we created genetically informed ecological niche models (gENMs) using Populus fremontii (Salicaceae), a widespread tree species in which prior common garden experiments demonstrate strong evidence for local adaptation. Four major findings emerged: (i) gENMs predicted population occurrences with up to 12‐fold greater accuracy than models without genetic information; (ii) tests of niche similarity revealed that three ecotypes, identified on the basis of neutral genetic markers and locally adapted populations, are associated with differences in climate; (iii) our forecasts indicate that ongoing climate change will likely shift these ecotypes further apart in geographic space, resulting in greater niche divergence; (iv) ecotypes that currently exhibit the largest geographic distribution and niche breadth appear to be buffered the most from climate change. As diverse agents of selection shape genetic variability and structure within species, we argue that gENMs will lead to more accurate predictions of species distributions under climate change.     

The shaping of genetic variation in edge‐of‐range populations under past and future climate change

With rates of climate change exceeding the rate at which many species are able to shift their range or adapt, it is important to understand how future changes are likely to affect biodiversity at all levels of organisation. Understanding past responses and extent of niche conservatism in climatic tolerance can help predict future consequences. We use an integrated approach to determine the genetic consequences of past and future climate changes on a bat species, Plecotus austriacus. Glacial refugia predicted by palaeo‐modelling match those identified from analyses of extant genetic diversity and model‐based inference of demographic history. Former refugial populations currently contain disproportionately high genetic diversity, but niche conservatism, shifts in suitable areas and barriers to migration mean that these hotspots of genetic diversity are under threat from future climate change. Evidence of population decline despite recent northward migration highlights the need to conserve leading‐edge populations for spearheading future range shifts.     

Phenological plasticity will not help all species adapt to climate change

Concerns are rising about the capacity of species to adapt quickly enough to climate change. In long‐lived organisms such as trees, genetic adaptation is slow, and how much phenotypic plasticity can help them cope with climate change remains largely unknown. Here, we assess whether, where and when phenological plasticity is and will be adaptive in three major European tree species. We use a process‐based species distribution model, parameterized with extensive ecological data, and manipulate plasticity to suppress phenological variations due to interannual, geographical and trend climate variability, under current and projected climatic conditions. We show that phenological plasticity is not always adaptive and mostly affects fitness at the margins of the species' distribution and climatic niche. Under current climatic conditions, phenological plasticity constrains the northern range limit of oak and beech and the southern range limit of pine. Under future climatic conditions, phenological plasticity becomes strongly adaptive towards the trailing edges of beech and oak, but severely constrains the range and niche of pine. Our results call for caution when interpreting geographical variation in trait means as adaptive, and strongly point towards species distribution models explicitly taking phenotypic plasticity into account when forecasting species distribution under climate change scenarios.     

Individual heterogeneity in life histories and eco‐evolutionary dynamics

Individual heterogeneity in life history shapes eco‐evolutionary processes, and unobserved heterogeneity can affect demographic outputs characterising life history and population dynamical properties. Demographic frameworks like matrix models or integral projection models represent powerful approaches to disentangle mechanisms linking individual life histories and population‐level processes. Recent developments have provided important steps towards their application to study eco‐evolutionary dynamics, but so far individual heterogeneity has largely been ignored. Here, we present a general demographic framework that incorporates individual heterogeneity in a flexible way, by separating static and dynamic traits (discrete or continuous). First, we apply the framework to derive the consequences of ignoring heterogeneity for a range of widely used demographic outputs. A general conclusion is that besides the long‐term growth rate lambda, all parameters can be affected. Second, we discuss how the framework can help advance current demographic models of eco‐evolutionary dynamics, by incorporating individual heterogeneity. For both applications numerical examples are provided, including an empirical example for pike. For instance, we demonstrate that predicted demographic responses to climate warming can be reversed by increased heritability. We discuss how applications of this demographic framework incorporating individual heterogeneity can help answer key biological questions that require a detailed understanding of eco‐evolutionary dynamics.     

Predicting global change impacts on plant species’ distributions: Future challenges

Given the rate of projected environmental change for the 21st century, urgent adaptation and mitigation measures are required to slow down the on-going erosion of biodiversity. Even though increasing evidence shows that recent human-induced environmental changes have already triggered species’ range shifts, changes in phenology and species’ extinctions, accurate projections of species’ responses to future environmental changes are more difficult to ascertain. This is problematic, since there is a growing awareness of the need to adopt proactive conservation planning measures using forecasts of species’ responses to future environmental changes.

There is a substantial body of literature describing and assessing the impacts of various scenarios of climate and land-use change on species’ distributions. Model predictions include a wide range of assumptions and limitations that are widely acknowledged but compromise their use for developing reliable adaptation and mitigation strategies for biodiversity. Indeed, amongst the most used models, few, if any, explicitly deal with migration processes, the dynamics of population at the “trailing edge” of shifting populations, species’ interactions and the interaction between the effects of climate and land-use.

In this review, we propose two main avenues to progress the understanding and prediction of the different processes occurring on the leading and trailing edge of the species’ distribution in response to any global change phenomena. Deliberately focusing on plant species, we first explore the different ways to incorporate species’ migration in the existing modelling approaches, given data and knowledge limitations and the dual effects of climate and land-use factors. Secondly, we explore the mechanisms and processes happening at the trailing edge of a shifting species’ distribution and how to implement them into a modelling approach. We finally conclude this review with clear guidelines on how such modelling improvements will benefit conservation strategies in a changing world.