Niche differentiation, reproductive interference, and range expansion.

Understanding species distributions and predicting future range shifts requires considering all relevant abiotic factors and biotic interactions. Resource competition has received the most attention, but reproductive interference is another widespread biotic interaction that could influence species ranges. Rubyspot damselflies (Hetaerina spp.) exhibit a biogeographic pattern consistent with the hypothesis that reproductive interference has limited range expansion. Here, we use ecological niche models to evaluate whether this pattern could have instead been caused by niche differentiation. We found evidence for climatic niche differentiation, but the species that encounters the least reproductive interference has one of the narrowest and most peripheral niches. These findings strengthen the case that reproductive interference has limited range expansion and also provide a counterexample to the idea that release from negative species interactions triggers niche expansion. We propose that release from reproductive interference enables species to expand in range while specializing on the habitats most suitable for breeding.

Niche conservatism and spread explain introgression between native and invasive fish.

Hybridisation can be an important driver of evolutionary change, but hybridisation with invasive species can have adverse effects on native biodiversity. While hybridisation has been documented across taxa, there is limited understanding of ecological factors promoting patterns of hybridisation and the spatial distribution of hybrid individuals. We combined the results of ecological niche modelling (ENM) and restriction site-associated DNA sequencing to test theories of niche conservatism and biotic resistance on the success of invasion, admixture, and extent of introgression between native and non-native fishes. We related Maxent predictions of habitat suitability based on the native ranges of invasive Eastern Banded Killifish (Fundulus diaphanus diaphanus Lesueur 1817) and native Western Banded Killifish (Fundulus diaphanus menona Jordan and Copeland 1877) to admixture indices of individual Banded Killifish. We found that Eastern Banded Killifish predominated at sites predicted as suitable from their ENM, consistent with niche conservatism. Admixed individuals were more common as Eastern Banded Killifish habitat suitability declined. We also found that Eastern Banded Killifish were most common at sites closest to the presumed source of this invasion, whereas the proportion of admixed individuals increased with distance from the source of invasion. Lastly, we found little evidence that habitat suitability for Western Banded Killifish provides biotic resistance from either displacement by, or admixture with, invasive Eastern Banded Killifish. Our study demonstrates that ENMs can inform conservation-relevant outcomes between native and invasive taxa while emphasising the importance of protecting isolated Western Banded Killifish populations from invasive conspecifics.

High climatic ancestral affinity between the lineages of the Leishmania vector Psathyromyia shannoni sensu stricto (Diptera: Phlebotominae).

Psathyromyia (Psathyromyia) shannoni sensu stricto (Dyar) is a vector of Leishmania parasite and the second sandfly of medical importance with a wide geographical but discontinuous distribution in America. Preliminary genetic structure analysis using a mitochondrial marker shows that the species integrated by at least four lineages could be the result of ecological adaptations to different environmental scenarios, but this hypothesis had never been proven. The aim of the present study was to analyse whether the genetic structure that detected Pa. shannoni ss. is associated with divergence or conservatism niche. Using Ecological Niche Models (ENMs) theory, we estimated the potential distribution for each genetic lineage, and then, we evaluated the equivalency niche for assessing whether climatic niche was more different than expected. The ENMs identify different suitable distribution areas but the same climatic or ecological conditions for the genetic lineages of Pa. shannoni (conservatism niche). Our findings allow us to speculate that other potential processes or events could be related to the genetic differentiation of Pa. shannoni. These studies are important because they allow us to identify the factors that could restrict the potential distribution of the different lineages whose vectorial competence is still unknown.

Selection of climate variables in ant species distribution models: case study in South Korea.

The selection of explanatory variables is important in modeling prediction of changes in species distribution in response to climate change. In this study, we evaluated the importance of variable selection in species distribution models. We compared two different types of models for predicting the distribution of ant species: temperature-only and both temperature and precipitation. Ants were collected at 343 forest sites across South Korea from 2006 through 2009. We used a generalized additive model (GAM) to predict the future distribution of 16 species that showed significant responses to changes in climatic factors (temperature and/or precipitation). Four types of GAMs were constructed: temperature, temperature with interaction of precipitation, temperature and precipitation without interaction, and temperature and precipitation with interaction. Most species displayed similar results between the temperatureonly and the temperature and precipitation models. The results for predicted changes in species richness were different from the temperature-only model. This indicates higher uncertainty in the prediction of species richness, which is obtained by combining the prediction results of distribution change for each species, than in the prediction of distribution change. The turnover rate of the ant assemblages was predicted to increase with decreases in temperature and increases in elevation, which was consistent with other studies. Finally, our results showed that the prediction of the distribution or diversity of organisms responding to climate change is uncertain because of the high variability of the model outputs induced by the variables used in the models.

Forecasting the future of Fall armyworm Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae) in India using ecological niche model.

The Fall armyworm, Spodoptera frugiperda is the most notorious invasive pest species on maize, recently reported in India. The continuous spread of Fall armyworms to new ecological niches raises global concern. The current study is the first in India to forecast the suitability of a habitat for S. frugiperda using a maximum entropy algorithm. Predictions were made based on an analysis of the relationship between 109 occurrence records of S. frugiperda and pertinent historical, current, and predicted climatic data for the study area. The model indicated that S. frugiperda could thrive in different habitats under the current environmental circumstances, particularly in the west and south Indian states like Maharashtra, Tamil Nadu, and Karnataka. The model predicted that areas with higher latitudes, particularly in Uttar Pradesh, Odisha, West Bengal, and some portions of Telangana, Rajasthan, Chhattisgarh, and Madhya Pradesh, as well as some tracts of northeastern states like Assam and Arunachal Pradesh, would have highly climate-suitable conditions for S. frugiperda to occur in the future. The average AUC value was 0.852, which indicates excellent accuracy of the prediction. A Jackknife test of variables indicated that isothermality with the highest gain value was determining the potential geographic distribution of S. frugiperda. Our results will be useful for serving as an early warning tool to guide decision-making and prevent further spread toward new areas in India.

Climate has contributed to population diversification of Daphnia galeata across Eurasia.

Climate is a fundamental abiotic factor that plays a key role in driving the evolution, distribution and population diversification of species. However, there have been few investigations of genomic signatures of adaptation to local climatic conditions in cladocerans. Here, we have provided the first high-quality chromosome-level genome assembly (~143 Mb, scaffold N50 12.6 Mb) of the waterflea, Daphnia galeata, and investigated genomic variation in 22 populations from Central Europe and Eastern China. Our ecological-niche models suggested that the historic distribution of D. galeata in Eurasia was significantly affected by Quaternary climate fluctuations. We detected pronounced genomic and morphometric divergences between European and Chinese D. galeata populations. Such divergences could be partly explained by genomic signatures of thermal adaptation to distinct climate regimes: a set of candidate single-nucleotide polymorphisms (SNPs) potentially associated with climate were detected. These SNPs were in genes significantly enriched in the Gene ontology terms "determination of adult lifespan" and "translation repressor activity", and especially, mthl5 and SOD1 involved in the IIS pathway, and EIF4EBP2 involved in the target of the rapamycin signalling pathway. Our study indicates that certain alleles might be associated with particular temperature regimes, playing a functional role in shaping the population structure of D. galeata at a large geographical scale. These results highlight the potential role of molecular variation in the response to climate variation, in the context of global climate change.

Mechanistic forecasts of species responses to climate change: The promise of biophysical ecology.

A core challenge in global change biology is to predict how species will respond to future environmental change and to manage these responses. To make such predictions and management actions robust to novel futures, we need to accurately characterize how organisms experience their environments and the biological mechanisms by which they respond. All organisms are thermodynamically connected to their environments through the exchange of heat and water at fine spatial and temporal scales and this exchange can be captured with biophysical models. Although mechanistic models based on biophysical ecology have a long history of development and application, their use in global change biology remains limited despite their enormous promise and increasingly accessible software. We contend that greater understanding and training in the theory and methods of biophysical ecology is vital to expand their application. Our review shows how biophysical models can be implemented to understand and predict climate change impacts on species' behavior, phenology, survival, distribution, and abundance. It also illustrates the types of outputs that can be generated, and the data inputs required for different implementations. Examples range from simple calculations of body temperature at a particular site and time, to more complex analyses of species' distribution limits based on projected energy and water balances, accounting for behavior and phenology. We outline challenges that currently limit the widespread application of biophysical models relating to data availability, training, and the lack of common software ecosystems. We also discuss progress and future developments that could allow these models to be applied to many species across large spatial extents and timeframes. Finally, we highlight how biophysical models are uniquely suited to solve global change biology problems that involve predicting and interpreting responses to environmental variability and extremes, multiple or shifting constraints, and novel abiotic or biotic environments.

Hydric effects on thermal tolerances influence climate vulnerability in a high-latitude beetle.

Species' thermal tolerances are used to estimate climate vulnerability, but few studies consider the role of the hydric environment in shaping thermal tolerances. As environments become hotter and drier, organisms often respond by limiting water loss to lower the risk of desiccation; however, reducing water loss may produce trade-offs that lower thermal tolerances if respiration becomes inhibited. Here, we measured the sensitivity of water loss rate and critical thermal maximum (CTmax ) to precipitation in nature and laboratory experiments that exposed click beetles (Coleoptera: Elateridae) to acute- and long-term humidity treatments. We also took advantage of their unique clicking behavior to characterize subcritical thermal tolerances. We found higher water loss rates in the dry acclimation treatment compared to the humid, and water loss rates were 3.2-fold higher for individuals that had experienced a recent precipitation event compared to individuals that had not. Acute humidity treatments did not affect CTmax , but precipitation indirectly affected CTmax through its effect on water loss rates. Contrary to our prediction, we found that CTmax was negatively associated with water loss rate, such that individuals with high water loss rate exhibited a lower CTmax . We then incorporated the observed variation of CTmax into a mechanistic niche model that coupled leaf and click beetle temperatures to predict climate vulnerability. The simulations indicated that indices of climate vulnerability can be sensitive to the effects of water loss physiology on thermal tolerances; moreover, exposure to temperatures above subcritical thermal thresholds is expected to increase by as much as 3.3-fold under future warming scenarios. The correlation between water loss rate and CTmax identifies the need to study thermal tolerances from a "whole-organism" perspective that considers relationships between physiological traits, and the population-level variation in CTmax driven by water loss rate complicates using this metric as a straightforward proxy of climate vulnerability.

Biodiversity conservation adaptation to climate change: Protecting the actors or the stage.

To be able to protect biodiversity in coming decades, conservation strategies need to consider what sites will be important for species not just today but also in the future. Different methods have been proposed to identify places that will be important for species in the future. Two of the most frequently used methods, ecological niche modeling and climate resilience, have distinct aims. The former focuses on identifying the suitable environmental conditions for species, thus protecting the "actor," namely, the species, whereas the latter seeks to safeguard the "stage," or the landscape in which species occur. We used the two methods to identify climate refugia for 258 forest vertebrates under short- and long-term climatic changes in a biodiversity hotspot, the Appalachian ecoregion of the United States. We also evaluated the spatial congruence of the two approaches for a possible conservation application, that of protecting 30% of the Appalachian region, in line with recent national and international policy recommendations. We detected weak positive correlations between resilience scores and baseline vertebrate richness, estimated with ecological niche models for historical (baseline) climatic conditions. The correlations were stronger for amphibians and mammals than for birds and reptiles. Under climate change scenarios, the correlations between estimated vertebrate richness and resilience were also weakly positive; a positive correlation was detected only for amphibians. Locations with estimated future gain of suitable climatic conditions for vertebrates showed low correlation with resilience. Overall, our results indicate that climate resilience and ecological niche modeling approaches capture different characteristics of projected distributional changes of Appalachian vertebrates. A climate resilience (the stage) approach could be more effective in safeguarding species with low dispersal abilities, whereas an ecological niche modeling (the actor) approach could be more suitable for species with long-distance dispersal capacity because they may be more broadly impacted by climate and less sensitive to geophysical features captured by a climate resilience approach.

Climate suitability modeling for Anastrepha suspensa (Diptera: Tephritidae): current and future invasion risk analysis.

The Caribbean fruit fly, Anastrepha suspensa (Lower, 1862) (Diptera: Tephritidae), is a pest of significant economic importance in Central America and Florida (USA). This study was carried out to examine the influence of climate change on the space-time distribution of A. suspensa on temporal and spatial scales. The CLIMEX software was used to model the current distribution and for climate change. The future distribution was performed using two global climate models (GCMs), CSIRO-Mk3.0 (CS) and MIROC-H (MR), under the emission scenarios (SRES) A2 and A1B for the years 2050, 2080, and 2100. The results indicate a low potential for global distribution of A. suspensa in all scenarios studied. However, tropical areas were identified with high climatic suitability for A. suspensa in South America, Central America, Africa, and Oceania until the end of the century. Projections of areas with climatic suitability for A. suspensa can provide helpful information to develop preventive strategies of phytosanitary management avoiding economic impacts with the introduction of the species.

Realized niche shift of an invasive widow spider: drivers and impacts of human activities.

BACKGROUND: Predicting invasiveness requires an understanding of the propensity of a given species to thrive in areas with novel ecological challenges. Evaluation of realized niche shift of an invasive species in its invasive range, detecting the main drivers of the realized niche shift, and predicting the potential distribution of the species can provide important information for the management of populations of invasive species and the conservation of biodiversity. The Australian redback spider, Latrodectus hasselti, is a widow spider that is native to Australia and established in Japan, New Zealand, and Southeast Asia. We used ecological niche models and ordinal comparisons in an integrative method to compare the realized niches of native and invasive populations of this spider species. We also assessed the impact of several climatic predictor variables and human activity on this niche shift. We hypothesized that human impact is important for successful establishment of this anthropophilic species, and that climatic predictor variables may determine suitable habitat and thus predict invasive ranges. RESULTS: Our models showed that L. hasselti distributions are positively influenced by human impact in both of the native and invasive ranges. Maximum temperature was the most important climatic variable in predictions of the distribution of native populations, while precipitation seasonality was the most important in predictions of invasive populations. The realized niche of L. hasselti in its invasive range differed from that in its native range, indicating possible realized niche shift. CONCLUSIONS: We infer that a preference for human-disturbed environments may underlie invasion and establishment in this spider species, as anthropogenic habitat modifications could provide shelters from unsuitable climatic conditions and extreme climatic stresses to the spiders. Because Australia and the countries in which the species is invasive have differing climates, differences in the availability of certain climatic conditions could have played a role in the realized niche shift of L. hasselti.

Thermal tolerance in Drosophila: Repercussions for distribution, community coexistence and responses to climate change.

Here we combined controlled experiments and field surveys to determine if estimates of heat tolerance predict distributional ranges and phenology of different Drosophila species in southern South America. We contrasted thermal death time curves, which consider both magnitude and duration of the challenge to estimate heat tolerance, against the thermal range where populations are viable based on field surveys in an 8-year longitudinal study. We observed a strong correspondence of the physiological limits, the thermal niche for population growth, and the geographic ranges across studied species, which suggests that the thermal biology of different species provides a common currency to understand how species will respond to warming temperatures both at a local level and throughout their distribution range. Our approach represents a novel analytical toolbox to anticipate how natural communities of ectothermic organisms will respond to global warming.

Analysis of potential drivers of spatial and temporal changes in anisakid larvae infection levels in European hake, Merluccius merluccius (L.), from the North-East Atlantic fishing grounds.

We analysed the spatial and temporal variability of Anisakis larvae infection in hake (Merluccius merluccius) from the North-East Atlantic from 1998 to 2020 and the potential drivers (i.e., environmental and host abundance) of such variation. The results showed that hake from separate sea areas in the North Atlantic have marked differences in temporal abundance levels. Hake larger than 60 cm were all parasitized in all ICES (International Council for the Exploration of the Sea) subareas 6, 7, and 8. The belly flaps were the most parasitized parts of the flesh, accounting for 92% of the total. Individuals of Anisakis simplex, Anisakis pegreffii, Anisakis spp. and a hybrid of Anisakis simplex × pegreffii were genetically identified, and Anisakis simplex as the most abundant (88-100%). An ecological niche model of Anisakis occurrence in fishes in the NE Atlantic was built to define the thermal optimum and environmental ranges for salinity, depth, chlorophyll concentration, and diffuse attenuation. The temporal variability of anisakid infection in fishes in the last two decades indicated an increase in the NE Atlantic at an annual rate of 31.7 nematodes per total number of specimens examined per year. This rise in infection levels could be triggered by the increase in intermediate host fish stocks, especially hake in the area.

Seasonal spatial dynamics of butterfly migration.

Understanding the seasonal movements of migratory species underpins ecological studies. Several hundred butterfly species show migratory behaviour, yet the spatial pattern of these migrations is poorly understood. We developed climatic niche models for 405 migratory butterfly species globally to estimate patterns of seasonal movement and the distribution of seasonal habitat suitability. We found strong seasonal variation in habitat suitability for most migratory butterflies with >75% of pixels within their distributions showing seasonal switching in predicted occupancy for 85% of species. The greatest rate of seasonal switching occurred in the tropics. Several species showed extreme range fluctuations between seasons, exceeding 10-fold for 53 species (13%) and more than 100-fold for nine species (2%), suggesting that such species may be at elevated extinction risk. Our results can be used to search for the ecological processes that underpin migration in insects, as well as to design conservation interventions for declining migratory insects.

Major population splits coincide with episodes of rapid climate change in a forest-dependent bird.

Climate change influences population demography by altering patterns of gene flow and reproductive isolation. Direct mutation rates offer the possibility for accurate dating on the within-species level but are currently only available for a handful of vertebrate species. Here, we use the first directly estimated mutation rate in birds to study the evolutionary history of pied flycatchers (Ficedula hypoleuca). Using a combination of demographic inference and species distribution modelling, we show that all major population splits in this forest-dependent system occurred during periods of increased climate instability and rapid global temperature change. We show that the divergent Spanish subspecies originated during the Eemian-Weichselian transition 115-104 thousand years ago (kya), and not during the last glacial maximum (26.5-19 kya), as previously suggested. The magnitude and rates of climate change during the glacial-interglacial transitions that preceded population splits in pied flycatchers were similar to, or exceeded, those predicted to occur in the course of the current, human-induced climate crisis. As such, our results provide a timely reminder of the strong impact that episodes of climate instability and rapid temperature changes can have on species' evolutionary trajectories, with important implications for the natural world in the Anthropocene.

Plastid phylogenomics and insights into the inter-mountain dispersal of the Eastern African giant senecios (Dendrosenecio, Asteraceae).

Giant senecios (Dendrosenecio, Asteraceae), endemic to the tropical mountains of Eastern Africa, are one of the most conspicuous alpine plant groups in the world. Although the group has received substantial attention from researchers, its infrageneric relationships are contentious, and the speciation history remains poorly understood. In this study, whole chloroplast genome sequences of 46 individuals were used to reconstruct the phylogeny of giant senecios using Maximum Likelihood and Bayesian Inference methods. The divergence times of this emblematic group were estimated using fossil-based calibrations. Additionally, the ancestral areas were inferred, and ecological niche modeling was used to predict their suitable habitats. Phylogenetic analyses yielded two robustly supported clades. One clade included taxa sampled from Tanzania, while the other clade included species from other regions. Giant senecios likely originated from the North of Tanzania approximately 2.3 million years ago (highest posterior density 95%; 0.77-4.40), then rapidly radiated into the Kenyan and Ugandan mountains within the last one million years. The potential routes of dispersal have been proposed based on the inferred ancestral areas, estimated time, and predicted past suitable niches. Plio-Pleistocene climate oscillations and orogeny instigated early divergence of the genus. Whereas in situ radiation of giant senecios was chiefly driven by multiple long-distance dispersal events followed by episodes of vicariance, and allopatric speciation (geographic and/or altitudinal).

Coexistence barriers confine the poleward range of a globally distributed plant.

In the study of factors shaping species' poleward range boundaries, climatic constraints are often assigned greater importance than biotic interactions such as competition. However, theory suggests competition can truncate a species' fundamental niche in harsh environments. We test this by challenging a mechanistic niche model - containing explicit competition terms - to predict the poleward range boundaries of two globally distributed, ecologically similar aquatic plant species. Mechanistic competition models accurately predicted the northern range limits of our study species, outperforming competition-free mechanistic models and matching the predictive ability of statistical niche models fit to occurrence records. Using the framework of modern coexistence theory, we found that relative nonlinearity in competitors' responses to temperature fluctuations maintains their coexistence boundary, highlighting the importance of this fluctuation-dependent mechanism. Our results support a more nuanced, interactive role of climate and competition in determining range boundaries, and illustrate a practical, process-based approach to understanding the determinants of range limits.

Spatial predictors of genomic and phenotypic variation differ in a lowland Middle American bird (Icterus gularis).

Spatial patterns of intraspecific variation are shaped by geographical distance among populations, historical changes in gene flow and interactions with local environments. Although these factors are not mutually exclusive and operate on both genomic and phenotypic variation, it is unclear how they affect these two axes of variation. We address this question by exploring the predictors of genomic and phenotypic divergence in Icterus gularis, a broadly distributed Middle American bird that exhibits marked geographical variation in body size across its range. We combined a comprehensive single nucleotide polymorphism and phenotypic data set to test whether genome-wide genetic and phenotypic differentiation are best explained by (i) isolation by distance, (ii) isolation by history or (iii) isolation by environment. We find that the pronounced genetic and phenotypic variation in I. gularis are only partially correlated and differ regarding spatial predictors. Whereas genomic variation is largely explained by historical barriers to gene flow, phenotypic diversity can be best predicted by contemporary environmental heterogeneity. Our genomic analyses reveal strong phylogeographical structure coinciding with the Chivela Pass at the Isthmus of Tehuantepec that was formed during the Pleistocene, when populations were isolated in north-south refugia. In contrast, we found a strong association between body size and environmental variables, such as temperature and precipitation. The relationship between body size and local climate is consistent with a pattern produced by either natural selection or environmental plasticity. Overall, these results provide empirical evidence for why phenotypic and genomic data are often in conflict in taxonomic and phylogeographical studies.

Greater topoclimatic control of above- versus below-ground communities.

Assessing the degree to which climate explains the spatial distributions of different taxonomic and functional groups is essential for anticipating the effects of climate change on ecosystems. Most effort so far has focused on above-ground organisms, which offer only a partial view on the response of biodiversity to environmental gradients. Here including both above- and below-ground organisms, we quantified the degree of topoclimatic control on the occurrence patterns of >1,500 taxa and phylotypes along a c. 3,000 m elevation gradient, by fitting species distribution models. Higher model performances for animals and plants than for soil microbes (fungi, bacteria and protists) suggest that the direct influence of topoclimate is stronger on above-ground species than on below-ground microorganisms. Accordingly, direct climate change effects are predicted to be stronger for above-ground than for below-ground taxa, whereas factors expressing local soil microclimate and geochemistry are likely more important to explain and forecast the occurrence patterns of soil microbiota. Detailed mapping and future scenarios of soil microclimate and microhabitats, together with comparative studies of interacting and ecologically dependent above- and below-ground biota, are thus needed to understand and realistically forecast the future distribution of ecosystems.

Multiple axes of ecological vulnerability to climate change.

Observed ecological responses to climate change are highly individualistic across species and locations, and understanding the drivers of this variability is essential for management and conservation efforts. While it is clear that differences in exposure, sensitivity, and adaptive capacity all contribute to heterogeneity in climate change vulnerability, predicting these features at macroecological scales remains a critical challenge. We explore multiple drivers of heterogeneous vulnerability across the distributions of 96 vegetation types of the ecologically diverse western US, using data on observed climate trends from 1948 to 2014 to highlight emerging patterns of change. We ask three novel questions about factors potentially shaping vulnerability across the region: (a) How does sensitivity to different climate variables vary geographically and across vegetation classes? (b) How do multivariate climate exposure patterns interact with these sensitivities to shape vulnerability patterns? (c) How different are these vulnerability patterns according to three widely implemented vulnerability paradigms-niche novelty (decline in modeled suitability), temporal novelty (standardized anomaly), and spatial novelty (inbound climate velocity)-each of which uses a distinct frame of reference to quantify climate departure? We propose that considering these three novelty paradigms in combination could help improve our understanding and prediction of heterogeneous climate change responses, and we discuss the distinct climate adaptation strategies connected with different combinations of high and low novelty across the three metrics. Our results reveal a diverse mosaic of climate change vulnerability signatures across the region's plant communities. Each of the above factors contributes strongly to this heterogeneity: climate variable sensitivity exhibits clear patterns across vegetation types, multivariate climate change data reveal highly diverse exposure signatures across locations, and the three novelty paradigms diverge widely in their climate change vulnerability predictions. Together, these results shed light on potential drivers of individualistic climate change responses and may help to inform effective management strategies.