Modelling and analysing evolution of dispersal in populations at expanding range boundaries

Abstract 1. Species would be expected to shift northwards in response to current climate warming, but many are failing to do so because of fragmentation of breeding habitats. Dispersal is important for colonisation and an individual‐based spatially explicit model was developed to investigate impacts of habitat availability on the evolution of dispersal in expanding populations. Model output was compared with field data from the speckled wood butterfly Pararge aegeria, which currently is expanding its range in Britain. 2. During range expansion, models simulated positive linear relationships between dispersal and distance from the seed location. This pattern was observed regardless of quantity (100% to 10% habitat availability) or distribution (random vs. gradient distribution) of habitat, although higher dispersal evolved at expanding range margins in landscapes with greater quantity of habitat and in gradient landscapes. Increased dispersal was no longer evident in any landscape once populations had reached equilibrium; dispersal values returned to those of seed populations. However, in landscapes with the least quantity of habitat, reduced dispersal (below that of seed populations) was observed at equilibrium. 3. Evolutionary changes in adult flight morphology were examined in six populations of P. aegeria along a transect from the distribution core to an expanding range margin in England (spanning a latitudinal distance of >200 km). Empirical data were in agreement with model output and showed increased dispersal ability (larger and broader thoraxes, smaller abdomens, higher wing aspect ratios) with increasing distance from the distribution core. Increased dispersal ability was evident in populations from areas colonised >30 years previously, although dispersal changes were generally evident only in females. 4. Evolutionary increases in dispersal ability in expanding populations may help species track future climate changes and counteract impacts of habitat fragmentation by promoting colonisation. However, at the highest levels of habitat loss, increased dispersal was less evident during expansion and reduced dispersal was observed at equilibrium indicating that, for many species, continued habitat fragmentation is likely to outweigh any benefits from dispersal.     

Evolutionary refugia and ecological refuges: key concepts for conserving Australian arid zone freshwater biodiversity under climate change

Refugia have been suggested as priority sites for conservation under climate change because of their ability to facilitate survival of biota under adverse conditions. Here, we review the likely role of refugial habitats in conserving freshwater biota in arid Australian aquatic systems where the major long‐term climatic influence has been aridification. We introduce a conceptual model that characterizes evolutionary refugia and ecological refuges based on our review of the attributes of aquatic habitats and freshwater taxa (fishes and aquatic invertebrates) in arid Australia. We also identify methods of recognizing likely future refugia and approaches to assessing the vulnerability of arid‐adapted freshwater biota to a warming and drying climate. Evolutionary refugia in arid areas are characterized as permanent, groundwater‐dependent habitats (subterranean aquifers and springs) supporting vicariant relicts and short‐range endemics. Ecological refuges can vary across space and time, depending on the dispersal abilities of aquatic taxa and the geographical proximity and hydrological connectivity of aquatic habitats. The most important are the perennial waterbodies (both groundwater and surface water fed) that support obligate aquatic organisms. These species will persist where suitable habitats are available and dispersal pathways are maintained. For very mobile species (invertebrates with an aerial dispersal phase) evolutionary refugia may also act as ecological refuges. Evolutionary refugia are likely future refugia because their water source (groundwater) is decoupled from local precipitation. However, their biota is extremely vulnerable to changes in local conditions because population extinction risks cannot be abated by the dispersal of individuals from other sites. Conservation planning must incorporate a high level of protection for aquifers that support refugial sites. Ecological refuges are vulnerable to changes in regional climate because they have little thermal or hydrological buffering. Accordingly, conservation planning must focus on maintaining meta‐population processes, especially through dynamic connectivity between aquatic habitats at a landscape scale.     

Effects of changing climate on aquatic habitat and connectivity for remnant populations of a wide‐ranging frog species in an arid landscape

Amphibian species persisting in isolated streams and wetlands in desert environments can be susceptible to low connectivity, genetic isolation, and climate changes. We evaluated the past (1900–1930), recent (1981–2010), and future (2071–2100) climate suitability of the arid Great Basin (USA) for the Columbia spotted frog (Rana luteiventris) and assessed whether changes in surface water may affect connectivity for remaining populations. We developed a predictive model of current climate suitability and used it to predict the historic and future distribution of suitable climates. We then modeled changes in surface water availability at each time period. Finally, we quantified connectivity among existing populations on the basis of hydrology and correlated it with interpopulation genetic distance. We found that the area of the Great Basin with suitable climate conditions has declined by approximately 49% over the last century and will likely continue to decline under future climate scenarios. Climate conditions at currently occupied locations have been relatively stable over the last century, which may explain persistence at these sites. However, future climates at these currently occupied locations are predicted to become warmer throughout the year and drier during the frog's activity period (May – September). Fall and winter precipitation may increase, but as rain instead of snow. Earlier runoff and lower summer base flows may reduce connectivity between neighboring populations, which is already limited. Many of these changes could have negative effects on remaining populations over the next 50–80 years, but milder winters, longer growing seasons, and wetter falls might positively affect survival and dispersal. Collectively, however, seasonal shifts in temperature, precipitation, and stream flow patterns could reduce habitat suitability and connectivity for frogs and possibly other aquatic species inhabiting streams in this arid region.     

Pleistocene climatic changes drive diversification across a tropical savanna

Spatial responses of species to past climate change depend on both intrinsic traits (climatic niche breadth, dispersal rates) and the scale of climatic fluctuations across the landscape. New capabilities in generating and analysing population genomic data, along with spatial modelling, have unleashed our capacity to infer how past climate changes have shaped populations, and by extension, complex communities. Combining these approaches, we uncover lineage diversity across four codistributed lizards from the Australian Monsoonal Tropics and explore how varying climatic tolerances interact with regional climate history to generate common vs. disparate responses to late Pleistocene change. We find more divergent spatial structuring and temporal demographic responses in the drier Kimberley region compared to the more mesic and consistently suitable Top End. We hypothesize that, in general, the effects of species’ traits on sensitivity to climate fluctuation will be more evident in climatically marginal regions. If true, this points to the need in climatically marginal areas to craft more species‐(or trait)‐specific strategies for persistence under future climate change.     

Future Climate Change Will Favour Non-Specialist Mammals in the (Sub)Arctics

Arctic and subarctic (i.e., [sub]arctic) ecosystems are predicted to be particularly susceptible to climate change. The area of tundra is expected to decrease and temperate climates will extend further north, affecting species inhabiting northern environments. Consequently, species at high latitudes should be especially susceptible to climate change, likely experiencing significant range contractions. Contrary to these expectations, our modelling of species distributions suggests that predicted climate change up to 2080 will favour most mammals presently inhabiting (sub)arctic Europe. Assuming full dispersal ability, most species will benefit from climate change, except for a few cold-climate specialists. However, most resident species will contract their ranges if they are not able to track their climatic niches, but no species is predicted to go extinct. If climate would change far beyond current predictions, however, species might disappear. The reason for the relative stability of mammalian presence might be that arctic regions have experienced large climatic shifts in the past, filtering out sensitive and range-restricted taxa. We also provide evidence that for most (sub)arctic mammals it is not climate change per se that will threaten them, but possible constraints on their dispersal ability and changes in community composition. Such impacts of future changes in species communities should receive more attention in literature.     

The effects of climate change on a mega‐diverse country: predicted shifts in mammalian species richness and turnover in continental Ecuador

Ecuador has some of the greatest biodiversity in the world, sheltering global biodiversity hotspots in lowland and mountain regions. Climate change will likely have a major effect on these regions, but the consequences for faunal diversity and conservation remain unclear. To address this issue, we used an ensemble of eight species distribution models to predict future shifts and identify areas of high changes in species richness and species turnover for 201 mammals. We projected the distributions using two different climate change scenarios at the 2050 horizon and contrasted two extreme dispersal scenarios (no dispersal vs. full dispersal). Our results showed extended distributional shifts all over the country. For most groups, our results predicted that the current diversity of mammals in Ecuador would decrease significantly under all climate change scenarios and dispersal assumptions. The Northern Andes and the Amazonian region would remain diversity hotspots but with a significant decrease in the number of species. All predictions, including the most conservative scenarios in terms of dispersal and climate change, predicted major changes in the distribution of mammalian species diversity in Ecuador. Primates might be the most severely affected because they would have fewer suitable areas, compared with other mammals. Our work emphasizes the need for sound conservation strategies in Ecuador to mitigate the effects of climate change     

Climate and land use changes will degrade the configuration of the landscape for titi monkeys in eastern Brazil

Land use changes have profound effects on populations of Neotropical primates, and ongoing climate change is expected to aggravate this scenario. The titi monkeys from eastern Brazil (Callicebus personatus group) have been particularly affected by this process, with four of the five species now allocated to threatened conservation status categories. Here, we estimate the changes in the distribution of these titi monkeys caused by changes in both climate and land use. We also use demographic‐based, functional landscape metrics to assess the magnitude of the change in landscape conditions for the distribution predicted for each species. We built species distribution models (SDMs) based on maximum entropy for current and future conditions (2070), allowing for different global circulation models and contrasting scenarios of glasshouse gas concentrations. We refined the SDMs using a high‐resolution map of habitat remnants. We then calculated habitat availability and connectivity based on home‐range size and the dispersal limitations of the individual, in the context of a predicted loss of 10% of forest cover in the future. The landscape configuration is predicted to be degraded for all species, regardless of the climatic settings. This include reductions in the total cover of forest remnants, patch size and functional connectivity. As the landscape configuration should deteriorate severely in the future for all species, the prevention of further loss of populations will only be achieved through habitat restoration and reconnection to counteract the negative effects for these and several other co‐occurring species.     

Pushing the Pace of Tree Species Migration

Plants and animals have responded to past climate changes by migrating with habitable environments, sometimes shifting the boundaries of their geographic ranges by tens of kilometers per year or more. Species migrating in response to present climate conditions, however, must contend with landscapes fragmented by anthropogenic disturbance. We consider this problem in the context of wind-dispersed tree species. Mechanisms of long-distance seed dispersal make these species capable of rapid migration rates. Models of species-front migration suggest that even tree species with the capacity for long-distance dispersal will be unable to keep pace with future spatial changes in temperature gradients, exclusive of habitat fragmentation effects. Here we present a numerical model that captures the salient dynamics of migration by long-distance dispersal for a generic tree species. We then use the model to explore the possible effects of assisted colonization within a fragmented landscape under a simulated tree-planting scheme. Our results suggest that an assisted-colonization program could accelerate species-front migration rates enough to match the speed of climate change, but such a program would involve an environmental-sustainability intervention at a massive scale.     

The fate of European breeding birds under climate,land‐use and dispersal scenarios

Many species have already shifted their distributions in response to recent climate change. Here, we aimed at predicting the future breeding distributions of European birds under climate, land‐use, and dispersal scenarios. We predicted current and future distributions of 409 species within an ensemble forecast framework using seven species distribution models (SDMs), five climate scenarios and three emission and land‐use scenarios. We then compared results from SDMs using climate‐only variables, habitat‐only variables or both climate and habitat variables. In order to account for a species’ dispersal abilities, we used natal dispersal estimates and developed a probabilistic method that produced a dispersal scenario intermediate between the null and full dispersal scenarios generally considered in such studies. We then compared results from all scenarios in terms of future predicted range changes, range shifts, and variations in species richness. Modeling accuracy was better with climate‐only variables than with habitat‐only variables, and better with both climate and habitat variables. Habitat models predicted smaller range shifts and smaller variations in range size and species richness than climate models. Using both climate and habitat variables, it was predicted that the range of 71% of the species would decrease by 2050, with a 335 km median shift. Predicted variations in species richness showed large decreases in the southern regions of Europe, as well as increases, mainly in Scandinavia and northern Russia. The partial dispersal scenario was significantly different from the full dispersal scenario for 25% of the species, resulting in the local reduction of the future predicted species richness of up to 10%. We concluded that the breeding range of most European birds will decrease in spite of dispersal abilities close to a full dispersal hypothesis, and that given the contrasted predictions obtained when modeling climate change only and land‐use change only, both scenarios must be taken into consideration.     

Do species’ traits predict recent shifts at expanding range edges?

Although some organisms have moved to higher elevations and latitudes in response to recent climate change, there is little consensus regarding the capacity of different species to track rapid climate change via range shifts. Understanding species' abilities to shift ranges has important implications for assessing extinction risk and predicting future community structure. At an expanding front, colonization rates are determined jointly by rates of reproduction and dispersal. In addition, establishment of viable populations requires that individuals find suitable resources in novel habitats. Thus, species with greater dispersal ability, reproductive rate and ecological generalization should be more likely to expand into new regions under climate change. Here, we assess current evidence for the relationship between leading-edge range shifts and species' traits. We found expected relationships for several datasets, including diet breadth in North American Passeriformes and egg-laying habitat in British Odonata. However, models generally had low explanatory power. Thus, even statistically and biologically meaningful relationships are unlikely to be of predictive utility for conservation and management. Trait-based range shift forecasts face several challenges, including quantifying relevant natural history variation across large numbers of species and coupling these data with extrinsic factors such as habitat fragmentation and availability.     

Climate change impacts on biodiversity in Switzerland: A review

A noticeable increase in mean temperature has already been observed in Switzerland and summer temperatures up to 4.8 K warmer are expected by 2090. This article reviews the observed impacts of climate change on biodiversity and considers some perspectives for the future at the national level.The following impacts are already evident for all considered taxonomic groups: elevation shifts of distribution towards mountain summits, spread of thermophilous species, colonisation by new species from warmer areas and phenological shifts. Additionally, in the driest areas, increasing droughts are affecting tree survival and fish species are suffering from warm temperatures in lowland regions. These observations are coherent with model projections, and future changes will probably follow the current trends.These changes will likely cause extinctions for alpine species (competition, loss of habitat) and lowland species (temperature or drought stress). In the very urbanised Swiss landscape, the high fragmentation of the natural ecosystems will hinder the dispersal of many species towards mountains. Moreover, disruptions in species interactions caused by individual migration rates or phenological shifts are likely to have consequences for biodiversity. Conversely, the inertia of the ecosystems (species longevity, restricted dispersal) and the local persistence of populations will probably result in lower extinction rates than expected with some models, at least in 21st century. It is thus very difficult to estimate the impact of climate change in terms of species extinctions. A greater recognition by society of the intrinsic value of biodiversity and of its importance for our existence will be essential to put in place effective mitigation measures and to safeguard a maximum number of native species.     

Warming increases the spread of an invasive thistle

Background

Global warming and shifted precipitation regimes increasingly affect species abundances and distributions worldwide. Despite a large literature on species'' physiological, phenological, growth, and reproductive responses to such climate change, dispersal is rarely examined. Our study aims to test whether the dispersal ability of a non-native, wind-dispersed plant species is affected by climate change, and to quantify the ramifications for future invasion spread rates.

Methodology/Principal Findings

We experimentally increased temperature and precipitation in a two-cohort, factorial field study (n = 80). We found an overwhelming warming effect on plant life history: warming not only improved emergence, survival, and reproduction of the thistle Carduus nutans, but also elevated plant height, which increased seed dispersal distances. Using spatial population models, we demonstrate that these empirical warming effects on demographic vital rates, and dispersal parameters, greatly exacerbate spatial spread. Predicted levels of elevated winter precipitation decreased seed production per capitulum, but this only slightly offset the warming effect on spread. Using a spread rate decomposition technique (c*-LTRE), we also found that plant height-mediated changes in dispersal contribute most to increased spread rate under climate change.

Conclusions/Significance

We found that both dispersal and spread of this wind-dispersed plant species were strongly impacted by climate change. Dispersal responses to climate change can improve, or diminish, a species'' ability to track climate change spatially, and should not be overlooked. Methods that combine both demographic and dispersal responses thus will be an invaluable complement to projections of suitable habitat under climate change.     

Range shift and loss of genetic diversity under climate change in Caryocar brasiliense, a Neotropical tree species

Many species have suffered reduction in habitable area due to recent climate change, but few studies evaluated how these range collapses will impact genetic diversity. Here, we modeled shifts in the species’ geographical range to evaluate how genetic diversity of Caryocar brasiliense will change as a consequence of predicted climate change in the next 50 years. A total of 135 records of species occurrence were obtained to model species’ distribution based on the current environment using MAXENT and forecasting future distribution using a combination of three coupled atmospheric–oceanic global circulation models. Genetic parameters were estimated based on the polymorphism at ten microsatellite loci for 466 individuals. Our results show that climatic suitable areas for C. brasiliense will be restricted to the southernmost distribution of savanna vegetation. Genetic diversity and the number of alleles may decrease slowly if populations persist in regions up to 0.5 of environmental suitability estimated by MAXENT, but will sharply decrease above this level. Nevertheless, deviation from mutation–drift equilibrium is significant even if a small amount of local populations is lost. More climatic suitable areas in the future will be in the most disturbed regions in Brazil, and populations that will persist there are those with higher levels of inbreeding at present. This may impose several threats to the species, including the limited capacity to cope with ongoing climatic changes by adaptation and constraints to dispersal.     

Projected changes in distributions of Australian tropical savanna birds under climate change using three dispersal scenarios

Identifying the species most vulnerable to extinction as a result of climate change is a necessary first step in mitigating biodiversity decline. Species distribution modeling (SDM) is a commonly used tool to assess potential climate change impacts on distributions of species. We use SDMs to predict geographic ranges for 243 birds of Australian tropical savannas, and to project changes in species richness and ranges under a future climate scenario between 1990 and 2080. Realistic predictions require recognition of the variability in species capacity to track climatically suitable environments. Here we assess the effect of dispersal on model results by using three approaches: full dispersal, no dispersal and a partial-dispersal scenario permitting species to track climate change at a rate of 30 km per decade. As expected, the projected distributions and richness patterns are highly sensitive to the dispersal scenario. Projected future range sizes decreased for 66% of species if full dispersal was assumed, but for 89% of species when no dispersal was assumed. However, realistic future predictions should not assume a single dispersal scenario for all species and as such, we assigned each species to the most appropriate dispersal category based on individual mobility and habitat specificity; this permitted the best estimates of where species will be in the future. Under this "realistic" dispersal scenario, projected ranges sizes decreased for 67% of species but showed that migratory and tropical-endemic birds are predicted to benefit from climate change with increasing distributional area. Richness hotspots of tropical savanna birds are expected to move, increasing in southern savannas and southward along the east coast of Australia, but decreasing in the arid zone. Understanding the complexity of effects of climate change on species' range sizes by incorporating dispersal capacities is a crucial step toward developing adaptation policies for the conservation of vulnerable species.     

Fine-scale genetic structure in a salamander with two reproductive modes: Does reproductive mode affect dispersal?

Reproduction is intimately linked with dispersal, but the effects of changes in reproductive strategies on dispersal have received little attention. Such changes have occurred in many taxonomic groups, resulting in profound alterations in life-history. In amphibians, many species shifted from oviparous/larviparous aquatic reproduction (deposition of eggs or pre-metamorphic larvae in water) to pueriparous terrestrial reproduction (parturition of terrestrial juveniles). The latter provides greater independence from water by skipping the aquatic larval stage; however, the eco-evolutionary implications of this evolutionary step have been underexplored, largely because reproductive modes rarely vary at the intraspecific level, preventing meaningful comparisons. We studied the effects of a transition to pueriparity on dispersal and fine-scale genetic structure in the fire salamander (Salamandra salamandra), a species exhibiting two co-occurring reproductive modes: larviparity and pueriparity. We performed genetic analyses (parentage and genetic spatial autocorrelation) using 11 microsatellite loci to compare dispersal and fine-scale genetic structure in three larviparous and three pueriparous populations (354 individuals in total). We did not find significant differences between reproductive modes, but in some larviparous populations movement patterns may be influenced by site-specific features (type of water bodies), possibly due to passive water-borne dispersal of larvae along streams. Additionally, females (especially larviparous ones) appeared to be more philopatric, while males showed greater variation in dispersal distances. This study also points to future avenues of research to better understand the eco-evolutionary implications of changes in reproductive modes in amphibians.     

Range shifts or extinction? Ancient DNA and distribution modelling reveal past and future responses to climate warming in cold‐adapted birds

Global warming is predicted to cause substantial habitat rearrangements, with the most severe effects expected to occur in high‐latitude biomes. However, one major uncertainty is whether species will be able to shift their ranges to keep pace with climate‐driven environmental changes. Many recent studies on mammals have shown that past range contractions have been associated with local extinctions rather than survival by habitat tracking. Here, we have used an interdisciplinary approach that combines ancient DNA techniques, coalescent simulations and species distribution modelling, to investigate how two common cold‐adapted bird species, willow and rock ptarmigan (Lagopus lagopus and Lagopus muta), respond to long‐term climate warming. Contrary to previous findings in mammals, we demonstrate a genetic continuity in Europe over the last 20 millennia. Results from back‐casted species distribution models suggest that this continuity may have been facilitated by uninterrupted habitat availability and potentially also the greater dispersal ability of birds. However, our predictions show that in the near future, some isolated regions will have little suitable habitat left, implying a future decrease in local populations at a scale unprecedented since the last glacial maximum.     

The importance of phenotypic plasticity and local adaptation in driving intraspecific variability in thermal niches of marine macrophytes

Climate change is driving the redistribution of species at a global scale and documenting and predicting species' responses to warming is a principal focus of contemporary ecology. When interpreting and predicting their responses to warming, species are generally treated as single homogenous physiological units. However, local adaptation and phenotypic plasticity can result in intraspecific differences in thermal niche. Therefore, population loss may also not only occur from trailing edges. In species with low dispersal capacity this will have profound impacts for the species as a whole, as local population loss will not be offset by immigration of warm tolerant individuals. Recent evidence from terrestrial forests has shown that incorporation of intraspecific variation in thermal niche is vital to accurately predicting species responses to warming. However, marine macrophytes (i.e. seagrasses and seaweeds) that form some of the world's most productive and diverse ecosystems have not been examined in the same context. We conducted a literature review to determine how common intraspecific variation in thermal physiology is in marine macrophytes. We find that 90% of studies identified (n = 42) found clear differences in thermal niche between geographically separated populations. Therefore, non‐trailing edge populations may also be vulnerable to future warming trends and given their limited dispersal capacity, such population loss may not be offset by immigration. We also explore how next generation sequencing (NGS) is allowing unprecedented mechanistic insight into plasticity and adaptation. We conclude that in the ‘genomic era’ it may be possible to link understanding of plasticity and adaptation at the genetic level through to changes in populations providing novel insights on the redistribution of populations under future climate change.     

The impacts of climate change on the annual cycles of birds

Organisms living today are descended from ancestors that experienced considerable climate change in the past. However, they are currently presented with many new, man-made challenges, including rapid climate change. Migration and reproduction of many avian species are controlled by endogenous mechanisms that have been under intense selection over time to ensure that arrival to and departure from breeding grounds is synchronized with moderate temperatures, peak food availability and availability of nesting sites. The timing of egg laying is determined, usually by both endogenous clocks and local factors, so that food availability is near optimal for raising young. Climate change is causing mismatches in food supplies, snow cover and other factors that could severely impact successful migration and reproduction of avian populations unless they are able to adjust to new conditions. Resident (non-migratory) birds also face challenges if precipitation and/or temperature patterns vary in ways that result in mismatches of food and breeding. Predictions that many existing climates will disappear and novel climates will appear in the future suggest that communities will be dramatically restructured by extinctions and changes in range distributions. Species that persist into future climates may be able to do so in part owing to the genetic heritage passed down from ancestors who survived climate changes in the past.     

Comparative landscape genetics reveals the evolution of viviparity reduces genetic connectivity in fire salamanders

Evolutionary changes in reproductive mode may affect co‐evolving traits, such as dispersal, although this subject remains largely underexplored. The shift from aquatic oviparous or larviparous reproduction to terrestrial viviparous reproduction in some amphibians entails skipping the aquatic larval stage and, thus, greater independence from water. Accordingly, amphibians exhibiting terrestrial viviparous reproduction may potentially disperse across a wider variety of suboptimal habitats and increase population connectivity in fragmented landscapes compared to aquatic‐breeding species. We investigated this hypothesis in the fire salamander (Salamandra salamandra), which exhibits both aquatic‐ (larviparity) and terrestrial‐breeding (viviparity) strategies. We genotyped 426 larviparous and 360 viviparous adult salamanders for 13 microsatellite loci and sequenced a mitochondrial marker for 133 larviparous and 119 viviparous individuals to compare population connectivity and landscape resistance to gene flow within a landscape genetics framework. Contrary to our predictions, viviparous populations exhibited greater differentiation and reduced genetic connectivity compared to larviparous populations. Landscape genetic analyses indicate viviparity may be partially responsible for this pattern, as water courses comprised a significant barrier only in viviparous salamanders, probably due to their fully terrestrial life cycle. Agricultural areas and, to a lesser extent, topography also decreased genetic connectivity in both larviparous and viviparous populations. This study is one of very few to explicitly demonstrate the evolution of a derived reproductive mode affects patterns of genetic connectivity. Our findings open avenues for future research to better understand the eco‐evolutionary implications underlying the emergence of terrestrial reproduction in amphibians.