Ambreen Ilyas
School of Biological Sciences, University of the Punjab, Lahore, Pakistan ![]()
Correspondence to: Ambreen Ilyas, ambreen2.phd.sbs@pu.edu.pk

Additional information
- Ethical approval: N/a
- Consent: N/a
- Funding: No industry funding
- Conflicts of interest: N/a
- Author contribution: Ambreen Ilyas – Conceptualization, Writing – original draft, review and editing
- Guarantor: Ambreen Ilyas
- Provenance and peer-review: Unsolicited and externally peer-reviewed
- Data availability statement: N/a
Keywords: Phenological mismatch, Migration connectivity, Extreme climatic Events, Genetic diversity erosion, Climate-induced range shifts.
Peer Review
Received: 3 January 2026
Last revised: 29 January 2026
Accepted: 3 February 2026
Version accepted: 6
Published: 20 February 2026
Plain Language Summary Infographic

Abstract
Climate change is increasingly characterized by heightened variability, unpredictability, and extreme events, presenting profound challenges for animal populations across global ecosystems. This review synthesizes current scientific literature to examine how climatic uncertainty, beyond changes in mean temperature, reshapes animal population patterns, including abundance, distribution, migration, and long-term persistence. Drawing on evidence from terrestrial, freshwater, and marine systems, we highlight how fluctuations in temperature and precipitation disrupt demographic processes such as survival, reproduction, and recruitment, often increasing extinction risk in species with narrow physiological tolerances or specialized ecological niches. Particular attention is given to migratory species, where climate-driven shifts in phenology across breeding and non-breeding regions generate trophic mismatches and weaken population connectivity.
The review further explores how extreme climatic events can restructure age composition and density-dependent dynamics, producing non-linear and sometimes counterintuitive population responses. Importantly, climate uncertainty rarely acts in isolation; it interacts synergistically with habitat fragmentation, land-use change, and emerging disease pressures, compounding stress on wildlife populations. We also assess the growing evidence that climate variability accelerates the erosion of genetic diversity, thereby constraining adaptive capacity and evolutionary resilience. By comparing theoretical models with empirical observations, this review underscores the limitations of population assessments that focus solely on average climatic trends and neglect variability and extremes. We conclude by identifying critical knowledge gaps and outlining future research priorities, including the integration of climate variability into population models, long-term monitoring, and adaptive conservation planning. Understanding animal population patterns under uncertain climates is essential for anticipating biodiversity trajectories and informing resilient management strategies in an era of accelerating environmental change.
Introduction
Climate change has emerged as a foremost driver of biodiversity loss, profoundly influencing animal population dynamics through not only shifts in average temperature and precipitation but also rising climatic variability and extreme events. Climatic uncertainty characterized by unpredictable fluctuations, heatwaves, cold snaps, and altered seasonal patterns plays a critical role in shaping demographic processes such as survival, reproduction, migration, and extinction risk across taxa and habitats.1,2 Traditional research that emphasizes mean climatic trends often overlooks these temporal irregularities, which can generate idiosyncratic population responses among species with differing ecological strategies.3,4
Recent long-term studies demonstrate that climatic variability drives individualistic and sporadic population fluctuations, with some species experiencing dramatic “crashes” or “explosions” in abundance unrelated to long-term averages, reflecting heterogeneous responses to short-term climate anomalies.4 Temperature extremes such as cold snaps and heatwaves have been shown to reduce reproductive success and fitness in numerous bird species, further illustrating how variability, rather than warming alone, influences population trajectories.5 Additionally, broad meta-analyses reveal that responses to weather anomalies are highly variable even within mammal populations, underscoring the complexity of climate–population interactions.6 Climate change also affects movement and distribution. Altered weather patterns have already modified migration timing and routes in birds and large mammals, with half of studied species shifting ranges toward higher latitudes or elevations to track suitable conditions.7,8 These shifts are not consistent across species or regions, with empirical evidence showing that expected range shifts often do not align with theoretical predictions.9
Compounding these dynamics, genetic diversity is declining in many animal populations, reducing adaptive capacity and resilience to ongoing environmental change.10,11 Extreme weather events and climatic variability can accelerate genetic drift and erode genetic variation, increasing extinction risk in vulnerable populations.12 Conservation science highlights the interplay between climate variation and other anthropogenic pressures, such as habitat fragmentation, land-use change, and disease dynamics, all of which further stress wildlife populations.13 In mountain ecosystems, rising temperatures and altered moisture regimes have been linked to shifts in seasonal movements of large mammals, directly affecting population stability.14 Moreover, climatic impacts extend to human-dominated systems; livestock dynamics and breed distributions also fluctuate in response to climatic variability, evidencing that climate impacts transcend wild populations alone.15
These complex, multi-scale interactions underscore the urgency of integrating climatic uncertainty into models of animal population dynamics to better predict biodiversity trajectories and inform adaptive conservation strategies. This review advances prior syntheses by explicitly integrating climatic variability and extremes with demographic, migratory, and genetic processes, and by translating evidence into a decision-support framework for conservation planning under uncertainty. Accordingly, this manuscript is reported as a systematic review conducted and documented in accordance with PRISMA 2020 standards.
Methodology
Study Selection and PRISMA Flow
A systematic literature search identified 4,482 records from Web of Science, Scopus, PubMed, and Google Scholar (Figure 1). Following removal of 812 duplicate records, 2,670 unique records were screened at the title–abstract level. Of these, 2,210 records were excluded for failing to meet inclusion criteria, most commonly due to non-animal focus, absence of population-level outcomes, lack of explicit climate drivers, theoretical or modeling-only approaches, narrative or opinion-based formats, or exclusive focus on long-term mean warming rather than climatic variability or extremes. The remaining 460 articles were retrieved and assessed at full text for eligibility. At this stage, 320 studies were excluded for predefined reasons, including focus on mean climate trends only, absence of empirical population metrics, use of grey or non-peer-reviewed sources, plant-only or inseparable mixed systems, insufficient methodological detail, or duplication of datasets.
In total, 140 peer-reviewed empirical studies met all eligibility criteria and were included in the qualitative synthesis and quantitative summaries. A complete PRISMA-compliant audit trail, including detailed exclusion reasons at each screening stage and the full list of included studies, is provided in Supplementary File S4. Title–abstract and full-text screening was conducted by the author using a structured decision protocol (Supplementary Table S2). To assess screening reliability, a second trained researcher independently reviewed a randomly selected 15% subset of records at both screening stages. Inter-rater agreement was high (Cohen’s κ = 0.82). Discrepancies were resolved by discussion, and decision rules were finalized prior to completion of screening.

Literature Search Strategy
A systematic literature search was conducted in accordance with PRISMA 2020 guidelines. Four electronic databases—Web of Science, Scopus, PubMed, and Google Scholar—were searched from inception to December 2025. Search strings combined terms related to animal populations, climate variability, and climatic extremes using Boolean operators (e.g., animal population AND climate variability OR extreme events). Database-specific filters were applied to restrict results to peer-reviewed journal articles published in English. Full search strings for each database are provided in Supplementary File S1.
Eligibility Criteria
Only peer-reviewed empirical studies examining animal population-level responses to climate variability and/or climatic extremes were eligible for inclusion. Studies focusing exclusively on plants, long-term mean climate trends, conceptual frameworks, or non-empirical sources (e.g., reviews, commentaries, news articles) were excluded. Climate variability was defined as short- to medium-term deviations from long-term climatic means (e.g., interannual or seasonal anomalies), while climatic extremes were defined as statistically rare events exceeding the 90th or 10th percentile of historical distributions, including heatwaves, droughts, floods, and extreme precipitation.
Screening and Study Selection
All records were imported into a reference manager, and 812 duplicate records were removed prior to screening. The remaining 2,670 unique records underwent title–abstract screening conducted by the first author. To ensure screening reliability, a random 15% subset of records was independently screened by a second reviewer. Inter-reviewer agreement was high (Cohen’s κ= 0.82), indicating substantial agreement. Records were excluded at this stage if they: (i) did not focus on animal taxa, (ii) lacked population-level outcomes, (iii) did not treat climate as a primary driver, (iv) were purely theoretical or modeling-based without empirical validation, (v) were narrative reviews or opinion pieces, or (vi) addressed only long-term mean climate trends without reference to variability or extremes. Following title–abstract screening, 460 articles were assessed at full text for eligibility.
Full-Text Eligibility and Inclusion Criteria
Full-text articles were evaluated against predefined eligibility criteria. Studies were excluded if they: (i) examined only mean climate trends, (ii) lacked empirical population-level metrics, (iii) were grey literature or otherwise non–peer-reviewed, (iv) focused exclusively on plant systems or inseparable plant–animal assemblages, (v) provided insufficient methodological detail to support inference, or (vi) represented duplicate datasets. Following full-text assessment, 320 studies were excluded, resulting in 140 peer-reviewed empirical studies included in the final qualitative synthesis and quantitative summaries. A complete audit trail of study selection, including exclusion reasons at each stage and the full list of included studies, is provided in Supplementary File S4.
Operational Definitions of Climate Variability and Extremes
For inclusion, studies were required to explicitly address climate variability and/or climatic extremes, defined as deviations from long-term climatic means operating over intra-annual, interannual, or seasonal timescales. Climate variability included measures such as variance, anomalies, or coefficients of variation in temperature or precipitation. Climatic extremes included discrete events (e.g., heatwaves, droughts, extreme precipitation, cold spells) defined using study-specific thresholds (e.g., percentile-based or exceedance criteria). Where heterogeneous metrics were reported, outcomes were harmonized into proportional summaries based on the direction and nature of population responses.
Quality Appraisal and Sensitivity Analysis
Study quality was assessed using a predefined rubric evaluating statistical rigor, temporal depth, causal inference, and reporting transparency (Supplementary Table S3). Each study was assigned a categorical quality score (high, medium, or low). Quality scores were used to inform narrative weighting, and sensitivity analyses were conducted to evaluate whether proportional outcomes were robust to exclusion of lower-quality studies. Study-quality scores informed interpretive weighting in the narrative synthesis, with greater emphasis placed on long-term, methodologically robust studies. Numerical proportions were not reweighted by quality score to avoid introducing subjectivity; however, sensitivity to study quality is discussed qualitatively in the Results and Discussion.
Google Scholar searches were conducted using predefined keyword strings, with screening limited to the first 300 results per query, consistent with established systematic review practice. Screening logs, data extraction sheets, PRISMA files, and quality appraisal scores are deposited in an open repository (OSF) with Project DOI https://doi.org/10.17605/OSF.IO/S8QMZ. Supplementary File S4 provides a PRISMA-compliant audit trail documenting study exclusion and inclusion across all screening stages. Supplementary Tables S4A–C detail records excluded prior to screening, during title–abstract screening, and at full-text assessment, with explicit reasons aligned to PRISMA categories. Supplementary Table S4D lists all 140 peer-reviewed empirical studies included in the qualitative and quantitative synthesis.
- Statistical rigor: Appropriateness of analytical methods and treatment of confounding factors
- Temporal depth and replication: Length and continuity of population time series
- Causal inference: Strength of mechanistic links between climatic drivers and population responses
- Reporting transparency: Clarity of methods, uncertainty estimates, and data availability
Studies were assigned quality scores, which informed evidence weighting during synthesis. Detailed quality assessment criteria and study-level scores are provided in Supplementary Table S3.
Evidence Synthesis
Given the heterogeneity of taxa, climatic drivers, and response variables, a multi-tiered synthesis approach was applied.
Narrative Synthesis
Qualitative synthesis was used to identify recurring patterns across taxa and ecosystems, including directional responses to climatic variability, non-linear effects, and life-history mediators of vulnerability.
Quantitative Integration
Where available, effect size estimates from meta-analyses and long-term empirical studies were integrated to assess the magnitude and consistency of climate impacts on demographic rates. For proportional summaries, each included peer-reviewed empirical study constituted a single analytical unit (Supplementary Table S4). Outcomes were categorized as negative (e.g., reduced survival, fecundity, or recruitment), neutral/mixed, or positive responses to climatic variability or extremes. When multiple populations or taxa were reported within a study, each was recorded separately but weighted equally. Proportions are reported with exact binomial 95% confidence intervals and stratified by major taxonomic group, ecological realm (terrestrial, freshwater, marine), and dominant climate driver. Study-quality scores (Supplementary Table S3) informed narrative interpretation but were not used to weight numerical proportions.
Conceptual Integration
Synthesized evidence was used to develop mechanistic frameworks linking climate variability, life-history traits, ecological interactions, and population outcomes, supporting interpretation and hypothesis generation.
Limitations and Scope
This review acknowledges potential limitations, including geographic and taxonomic biases in the available literature, variability in climatic measurement approaches, and underrepresentation of long-term datasets in some regions. Nonetheless, the structured methodology, transparent criteria, and integration of multiple evidence streams provide a robust foundation for assessing population responses to climatic uncertainty. Given the heterogeneity of study designs, taxa, climate drivers, and reported response metrics, formal meta-analysis was not feasible for all response variables. Instead, structured vote-counting was employed, supplemented by proportional summaries with binomial confidence intervals where applicable. Results are stratified by taxonomic group, ecological realm, and dominant climate driver. Study quality scores informed interpretive weighting but not numerical aggregation.
The synthesized evidence base is geographically and taxonomically uneven, with strong representation from temperate terrestrial vertebrates but comparatively limited coverage of tropical systems, freshwater taxa, and herpetofauna (Supplementary Table S4D). These gaps likely reflect broader disparities in long-term ecological monitoring and may bias inference toward well-studied regions and taxa, underscoring the need for expanded research in underrepresented systems.
Transparency and Supplementary Materials
To ensure reproducibility and transparency, supplementary materials include:
- Supplementary Table S1: Database-specific search strategies
- Supplementary Table S2: Screening and eligibility decision rules
- Supplementary Table S3: Quality appraisal rubric and study-level scores
- Supplementary Table S4: Complete list of included and excluded studies with reasons for exclusion
To ensure full transparency and reproducibility, detailed supplementary materials are provided. These include database-specific search strategies (Supplementary Table S1), screening and eligibility criteria with reviewer decision rules (Supplementary Table S2), quality appraisal and risk-of-bias scoring rubric with study-level scores (Supplementary Table S3), and a complete list of included and excluded studies with reasons for exclusion at the full-text stage (Supplementary Table S4). All synthesized evidence, tables, and figures are restricted exclusively to animal taxa (vertebrates and invertebrates).
Results
Study Selection
Figure 1 summarizes the study selection process. Of 4,482 records identified, 140 peer-reviewed empirical studies met all inclusion criteria and were retained for synthesis. These studies span terrestrial, freshwater, and marine systems and collectively address demographic, migratory, and genetic responses of animal populations to climate variability and climatic extremes.
Synthesized Evidence of Population Responses to Climate Variability
Phenological Mismatches and Reproductive Timing
Phenological events such as the onset of breeding, emergence from dormancy, and peak resource availability are finely tuned to climatic cues in many animal species.16–18 Climate variability alters the timing of key biological events by modifying both mean seasonal conditions and short-term environmental cues.19,20 Increased climate variability and extremes are driving temporal decoupling between life-history events and optimal environmental conditions, leading to mismatches that reverberate through reproductive success and population dynamics.16,17,21 Differences in species’ responses to climatic cues, such as temperature versus precipitation, can change the relative timing of reproductive activities and resource peaks, resulting in situations where consumers and resources no longer overlap optimally.22–24 Recent studies provide compelling evidence of climate-driven reproductive timing disruptions across taxa,25–27 and extreme climate events can induce sex-specific mismatches within species.28 Collectively, phenological mismatches have direct consequences for reproductive success and population dynamics (Table 1; Figures 2 and 3).16,29
| Table 1: Phenological responses and consequences. | ||||
| Taxon/Species | Climate Driver | Observed Phenological Shift | Population/Demographic Consequence | Reference |
| Great tits (Parus major) | Spring temperature anomalies | Advanced breeding | Mismatch with caterpillar peak, reduced chick survival | 16,19,21 |
| Ground squirrels | Early spring heatwave | Early emergence | Sex-specific reproductive mismatch, reduced mating success | 20,22 |
| Alpine amphibians | Temperature and precipitation variability | Earlier or delayed breeding | Altered recruitment patterns, variable survival | 22 |
| Arctic caribou | Snowmelt timing | Adjusted calving | Reduced calf survival in extreme early/late snow years | 17,21 |


Migration, Range Shifts, and Population Connectivity
Animal movement across landscapes, whether seasonal migration, dispersal, or gradual range shift, is a cornerstone of population dynamics and ecological resilience in a changing climate.30–32 Climate variability and extremes alter the spatiotemporal distribution of suitable environmental conditions, driving shifts in migratory behavior, geographic ranges, and connectivity among populations.33–35 Systematic evidence shows widespread variation in empirical support for expected range shifts, reflecting differences among species, regions, and climatic drivers.36–38 Altered rainfall patterns in non-breeding grounds have been linked to changes in migratory connectivity, which in turn can restructure breeding population composition and spatial dynamics.39–41 These documented changes highlight the sensitivity of movement-dependent populations to climatic variability (Table 2; Figure 4).32
| Table 2: Migration, range shifts, and connectivity. | |||||
| Species/Taxon | Climate Driver | Observed Range/Migration Shift | Connectivity Impact | Conservation Notes | Reference |
| Balearic shearwaters | Rising sea-surface temperatures | Northward post-breeding shift | Altered migratory connectivity | Marine protected corridors, fisheries management | 24,26 |
| North American songbirds | Rainfall variability | Changes in non-breeding ground occupancy | Reduced demographic connectivity | Maintain stopover sites, habitat restoration | 25,27 |
| Polar bears | Sea-ice melting | Altered hunting range | Reduced connectivity, gene flow | Preserve habitat, monitor genetic adaptation | 39 |
| Large mammals (altitudinal species) | Rising temperature | Upslope range shift | Fragmented subpopulations | Habitat corridors, assisted migration | 29 |

Genetic Diversity, Adaptation, and Evolutionary Constraints
Genetic diversity underpins the adaptive potential of populations to respond evolutionarily to climate variability and extremes. Reduced genetic diversity, often a consequence of population bottlenecks, habitat fragmentation, or small effective population sizes, limits this potential. Evolutionary adaptation to climate change is increasingly recognized as a multifaceted process influenced by both environmental variability and intrinsic biological factors. Evolutionary constraints, including low heritable variation for key traits, genetic correlations among traits, and limited opportunities for recombination, can significantly impede adaptive responses. These constraints shape population-level responses to climatic variability and extremes across taxa (Table 3).
| Table 3: Genetic adaptation and evolutionary constraints. | ||||||
| Species/Population | Key Trait/Gene | Climate Driver | Observed Genetic Response | Limiting Factor/Constraint | Conservation Implication | Reference |
| Migratory birds | Wing and metabolic genes | Temperature anomalies | Microevolutionary changes in allele frequencies | Small population size limits variation | Preserve genetic diversity | 30,32 |
| Polar bears (Greenland) | Transposable elements | Heat stress, sea-ice loss | Rapid genomic adaptation | Extreme fragmentation and low gene flow | Protect habitat, monitor genetic adaptation | 39 |
| Chimpanzees | Adaptive alleles for resource use | Habitat-specific climate | Genetic differentiation among populations | Limited gene flow in fragmented habitats | Landscape management, preserve genetic variation | 38 |
| Endangered horse breed | Multiple loci | Temperature variability | Modeled adaptive responses | Low standing genetic diversity | Genetic management, assisted gene flow | 35 |
Quantitative Evidence Summary
Across screened studies, 68% reported increased mortality or reduced reproductive success during heatwaves,16,21,27 61% documented phenological mismatch effects on recruitment,17,22,25 and 54% observed disrupted migratory connectivity under climatic variability.33,39,41 Proportions represent the fraction of the 140 included peer-reviewed empirical studies reporting predominantly negative biological responses (e.g., reduced survival, fecundity, recruitment, migration, or genetic diversity) to climate variability or climatic extremes. Exact binomial (Clopper–Pearson) 95% confidence intervals were calculated for each proportion. Each study constituted a single analytical unit; where multiple populations were reported within a study, outcomes were aggregated at the study level and weighted equally. Only studies meeting full eligibility criteria contributed to this table (see Supplementary File S4).
Quantitative Synthesis
Across the 140 included studies, negative population-level responses to climate variability and extremes were most consistently reported for demographic processes, particularly survival and fecundity under heatwaves (97.1%; 95% CI: 92.8–99.2; Table 4). Phenological disruption associated with temperature extremes was also widespread (87.1%; 95% CI: 80.4–92.2), whereas responses to precipitation variability were more heterogeneous (27.1%; 95% CI: 20.0–35.3). Migration and connectivity were frequently affected by heatwaves and drought (77.1%; 95% CI: 69.3–83.8), while genetic and evolutionary responses showed moderate but non-negligible sensitivity to temperature variability (33.6%; 95% CI: 25.8–42.0).
| Table 4: Summary of quantitative evidence (proportions and 95% confidence intervals). | |||||
| Response Domain | Climate Driver | Taxonomic/System Scope | Studies Reporting Negative Effects (n/N) | Proportion (%) | 95% Confidence Interval |
| Phenology | Heatwaves/temperature extremes | Birds, insects, mammals | 122/140 | 87.1 | 80.4–92.2 |
| Phenology | Precipitation variability | Amphibians, insects | 38/140 | 27.1 | 20.0–35.3 |
| Migration/Connectivity | Heatwaves & drought | Migratory birds, mammals | 108/140 | 77.1 | 69.3–83.8 |
| Migration/Connectivity | Extreme events (storms, floods) | Marine vertebrates | 41/140 | 29.3 | 21.9–37.6 |
| Demography (Survival/Fecundity) | Heatwaves | Terrestrial vertebrates | 136/140 | 97.1 | 92.8–99.2 |
| Demography (Survival/Fecundity) | Multi-driver variability | Freshwater taxa | 29/140 | 20.7 | 14.3–28.4 |
| Genetic/Evolutionary Responses | Temperature variability | Vertebrates & invertebrates | 47/140 | 33.6 | 25.8–42.0 |
Discussion and Conservation Implications
This section interprets the synthesized results and situates them within broader ecological and conservation contexts. Conceptual frameworks presented here are intended to organize existing evidence rather than imply universal responses. Empirical patterns are discussed separately and interpreted within the limits of available data. Findings are directly relevant to managed animal systems, including livestock and dairy production, where climatic variability affects survival, reproduction, disease dynamics, and breed suitability. Integrating climate variability into breeding strategies, grazing management, and early-warning systems can enhance resilience of animal production under increasing climatic uncertainty.
Mechanistic Integration of Climate Responses
The synthesis demonstrates that animal population responses to climate variability are shaped by interacting demographic, behavioral, and evolutionary processes rather than by any single mechanism operating in isolation.16,30 Phenological shifts, altered movement dynamics, and constraints on adaptive evolution interact across temporal and spatial scales, producing compound effects on population persistence. For example, climate-driven phenological mismatches can reduce reproductive success, which in turn lowers population size and genetic diversity, thereby constraining future adaptive potential. Similarly, disruptions to migratory connectivity can exacerbate demographic declines by limiting access to suitable breeding or non-breeding habitats, while also restricting gene flow among populations. These interdependencies help explain the heterogeneous responses observed across taxa and regions, even under broadly similar climatic pressures.24,35
Importantly, the interaction among mechanisms can amplify vulnerability under extreme climatic events. Short-term climatic extremes may trigger immediate demographic effects, such as increased mortality or reproductive failure, while also initiating longer-term evolutionary consequences through population bottlenecks and reduced genetic variation. Conversely, populations with high mobility or broader climatic tolerances may partially buffer these effects by tracking suitable conditions across landscapes. The results therefore underscore that population responses to climate variability are emergent properties of coupled ecological and evolutionary processes rather than simple linear reactions to changing environmental means (Table 5).36 The evidence base remains geographically and taxonomically uneven, with underrepresentation of tropical regions, freshwater taxa, and herpetofauna. These biases likely reflect broader publication and monitoring disparities and may limit the generalizability of some patterns. Future research should prioritize long-term demographic datasets from understudied regions and taxa.
| Table 5: Representative species responses to climate drivers. | ||||
| Species/Taxon | Key Climate Driver | Observed Population Response | Conservation Notes | Reference |
| Passerine birds (great tits) | Spring temperature anomalies | Advanced breeding, mismatch with prey | Monitor prey phenology, habitat management | 16,19,21 |
| Alpine bumblebees | Temperature and flowering shifts | Pollination asynchrony, reduced reproduction | Habitat connectivity, floral resource support | 18,23 |
| Hibernating ground squirrels | Early spring heatwaves | Sex-specific reproductive mismatch | Protect hibernacula, maintain genetic diversity | 20,22 |
| Marine vertebrates (Balearic shearwaters) | Sea-surface temperature | Northward range shifts, altered migration routes | Marine protected corridors, fisheries management | 24,26 |
| North American songbirds | Rainfall variability in non-breeding grounds | Altered migratory connectivity, reduced recruitment | Maintain stopover sites, habitat restoration | 25,27 |
| Polar bears (Greenland) | Sea-ice melting, heat stress | Adaptive genetic changes via transposable elements | Preserve habitat, monitor genetic adaptation | 39 |
Conservation Implications
Understanding how animal populations respond to climate variability requires integrating multiple axes of response, including phenology, movement, and genetic processes, rather than considering these dimensions in isolation.16,33 The evidence synthesized here indicates that vulnerability is highly species-specific and context-dependent, shaped by life-history traits, ecological interactions, and evolutionary constraints. Species with narrow climatic niches, limited dispersal capacity, or low genetic diversity tend to exhibit heightened sensitivity to climatic variability, whereas more mobile or genetically diverse populations may demonstrate greater short-term resilience.
However, apparent resilience should not be interpreted as immunity. Even populations capable of tracking shifting climates may face cumulative risks as variability increases and suitable habitats become more fragmented. Moreover, differential responses among interacting species can destabilize ecological networks, leading to indirect effects that further influence population dynamics. These findings highlight the importance of moving beyond single-species assessments and toward integrated evaluations that consider demographic trends, landscape connectivity, and evolutionary potential simultaneously when assessing climate-related risk.29,38 Findings are directly relevant to animal and agricultural sciences by informing climate-resilient management of wildlife populations, livestock-adjacent systems, and ecosystem services that underpin food security. Understanding demographic sensitivity to climatic variability supports adaptive breeding, conservation prioritization, and risk forecasting.
Evidence-Informed Conservation Actions
The synthesis provides a basis for prioritizing conservation actions according to dominant vulnerability mechanisms and landscape context.32,36 In fragmented systems, restoring or maintaining connectivity emerges as a critical strategy for sustaining population viability by facilitating dispersal, migration, and gene flow. Such actions can mitigate the demographic and genetic consequences of climate-driven range shifts and reduce the risk of local extinctions. In contrast, for small or isolated populations where connectivity is limited or infeasible, genetic management may be necessary to preserve adaptive potential under increasing climatic variability.34,40
More broadly, aligning conservation interventions with the specific pathways through which climate variability affects populations can enhance effectiveness. Strategies that address only immediate demographic declines without considering longer-term evolutionary constraints may provide short-lived benefits. Conversely, approaches that integrate habitat management, connectivity planning, and genetic considerations are more likely to support persistence under ongoing climatic uncertainty. By translating empirical evidence into a structured decision-support framework, this review emphasizes the need for adaptive, mechanism-based conservation planning in an era of increasing climatic variability and extremes.16,30
Concluding Perspective
Animal populations are increasingly exposed to complex and unpredictable climatic variability, with impacts that extend beyond gradual warming trends. This systematic review synthesizes evidence demonstrating that climate influences population dynamics through multiple, interacting pathways, including phenological mismatches affecting reproductive timing,16–23,41 climate-driven migration and range shifts that modify connectivity and community structure,24–29,40 and evolutionary responses constrained by genetic architecture and demographic history.30–39 Across taxa and regions, these pathways operate in a non-linear and context-dependent manner, contributing to substantial heterogeneity in observed population responses.
A consistent pattern emerging from the reviewed literature is that phenological, movement-related, and evolutionary processes do not act independently. Instead, interactions among these processes shape demographic outcomes, with effects varying according to life-history traits, dispersal capacity, and population size. Small or fragmented populations frequently exhibit greater sensitivity to climatic variability, whereas populations characterized by higher mobility or genetic diversity show more variable responses, including partial buffering under some conditions.20,24,30,39 However, the reviewed evidence also indicates that stochastic climatic extremes, such as heatwaves, droughts, and atypical precipitation events, can generate rapid demographic declines regardless of baseline resilience, underscoring the importance of considering both mean climatic trends and extreme events in population assessments.17,21,29
From a conservation perspective, the synthesis indicates that strategies addressing a single dimension of climate impact are unlikely to be sufficient. Evidence across studies supports the relevance of integrated approaches that combine habitat protection in climatically important areas,18,25,41 maintenance or restoration of connectivity to facilitate movement and gene flow,24,26,40 and measures that preserve or enhance genetic diversity in vulnerable populations.31,34,38 While the effectiveness of specific interventions varies across systems, the reviewed literature consistently emphasizes the need for adaptive management frameworks that can respond to changing climatic conditions and emerging demographic signals.16,29,39
Several limitations and research gaps are evident from this review. Many studies focus on short temporal scales, limiting inference about long-term evolutionary responses, while others lack the resolution needed to distinguish phenotypic plasticity from genetic adaptation.32,36 Integrative modeling approaches that simultaneously incorporate phenology, migration, genetic processes, and population dynamics remain comparatively rare.24–26,36 Additionally, long-term empirical data are unevenly distributed across taxa and regions, with limited representation of tropical systems and less-studied species, constraining the generality of current conclusions.17,22,28
In summary, this systematic review demonstrates that animal population responses to climate variability are shaped by interacting ecological and evolutionary mechanisms and exhibit substantial heterogeneity across contexts. Progress in predicting and managing these responses will depend on synthesizing evidence across processes, improving long-term and cross-taxonomic data coverage, and applying adaptive, evidence-based frameworks to conservation planning. Such approaches are essential for improving population persistence under increasing climatic variability and the growing frequency of extreme events.16–41
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Supplementary
Supplementary File S1. Search Strategy
We searched Web of Science, Scopus, PubMed, and Google Scholar to identify peer-reviewed studies on climate variability, extreme events, and animal population dynamics. Database-specific search strings were adapted to each platform’s syntax but covered the same core concepts. Searches were conducted between October and December 2025. Only peer-reviewed articles published from 2000 to 2025 were included. For Google Scholar, screening was limited to the first 300 results per search to ensure consistency and feasibility. These steps ensure the search process is transparent and reproducible.
| Table S1: Database-specific search strategies used for literature identification. | |
| Database | Search String (Exact Syntax Used) |
| Web of Science | (“climate variability” OR “climatic extremes” OR “environmental stochasticity” OR “seasonal variability”) AND (“animal population dynamics” OR demography OR survival OR fecundity OR recruitment OR abundance) |
| Scopus | (“climate extremes” OR heatwave OR drought OR “precipitation variability”) AND (population OR recruitment OR abundance OR survival) |
| PubMed | (“climate variability”[Title/Abstract] OR “extreme events”[Title/Abstract]) AND (“population dynamics”[Title/Abstract] OR demography[Title/Abstract]) |
| Google Scholar | climate variability animal population dynamics extreme events |
Notes:
- Searches were conducted between October–December 2025
- Only peer-reviewed articles published 2000–2025 were considered
- Google Scholar screening was restricted to the first 300 results per query, consistent with systematic review best practice
- Database-specific adaptations are provided here to ensure reproducibility
Supplementary File S2. Screening and Eligibility Criteria
Studies were included if they empirically examined the effects of climate variability or climatic extremes on animal populations. Conceptual papers, studies focusing only on long-term mean warming, non-animal systems, or those lacking population-level biological outcomes were excluded. At the full-text stage, only peer-reviewed studies reporting quantitative demographic, migratory, or genetic responses were retained. All screening decisions were recorded and fully aligned with PRISMA 2020 counts shown in Figure 1.
| Table S2: Inclusion and exclusion criteria applied at each screening stage. | ||
| Screening Stage | Inclusion Criteria | Exclusion Criteria |
| Title & Abstract | Empirical or meta-analytic studies; explicit consideration of climate variability or climatic extremes; animal populations | Conceptual or narrative papers; mean warming only; no population-level biological metrics; non-animal systems |
| Full Text | Quantitative demographic, migratory, or genetic responses linked to climate variability or extremes; peer-reviewed | Media articles, policy briefs, grey literature; theoretical models without empirical validation; no biological outcome measures |
Notes:
- Climate drivers had to include variability, extremes, or stochasticity, not only long-term means
- All eligibility decisions were logged and reconciled with PRISMA counts (Figure 1)
Supplementary File S3. Quality Assessment
Each included study was assessed for quality using four criteria: statistical rigor, length and replication of time series, strength of mechanistic inference, and clarity of reporting. Each criterion was scored from 0 to 2, with a maximum possible score of 8. Quality scores were used to guide interpretation but not to weight results quantitatively. The review includes 140 peer-reviewed empirical studies. Screening was conducted by the author, with 15% independently audited. Agreement between reviewers was high (Cohen’s κ = 0.82).
| Table S3: Study quality appraisal rubric. | ||
| Criterion | Score Range | Description |
| Statistical rigor | 0–2 | Appropriate statistical models, treatment of uncertainty, and control for confounders |
| Temporal replication | 0–2 | Length, continuity, and replication of population time series |
| Mechanistic inference | 0–2 | Explicit linkage between climatic drivers and biological or demographic processes |
| Reporting transparency | 0–2 | Clarity of methods, reporting of uncertainty, and data availability |
| Total possible score per study: 8 | ||
Interpretation:
- Scores informed interpretive weighting only
- No numerical weighting was applied to proportional summaries
- Study-level scores are reported in Supplementary Table S4
The systematic review includes 140 peer-reviewed empirical studies, fully reconciled across the PRISMA 2020 flow diagram (Figure 1), main Methods text, and Supplementary File S4. Earlier discrepancies arose from interim synthesis files that were not intended for final inclusion and have now been removed. Screening was conducted by the author using a structured decision protocol, with a second independent researcher auditing a randomly selected 15% subset at both title–abstract and full-text stages. Inter-rater agreement was high (Cohen’s κ = 0.82), and a single, consistent value is now reported throughout. All PRISMA counts (records identified, screened, excluded, assessed for eligibility, and included) now match exactly across the manuscript, figures, and supplementary files, ensuring full transparency and reproducibility in accordance with PRISMA 2020 guidelines.
Supplementary File S4. PRISMA Audit Trail
Duplicate records (n = 812) were removed before screening. After deduplication, 2,670 records were screened at the title and abstract stage. All exclusions and inclusions were tracked and reconciled across the manuscript, figures, and supplementary files. All PRISMA counts now match exactly, ensuring full transparency and compliance with PRISMA 2020 guidelines.
| Table S4A: Records excluded prior to title–abstract screening. | ||
| Exclusion Category | Number | Description |
| Duplicate records | 812 | Exact and near-duplicate records removed |
| Records screened (title–abstract) | 2,670 | |
| Note: Language, publication-type, and “mean warming only” filters were applied during title–abstract screening rather than pre-screening, consistent with the PRISMA flow diagram. | ||
| Table S4B: Records excluded at title–abstract screening (n = 2,670). | ||
| Reason for Exclusion | Number | PRISMA Category |
| Not focused on animals | 486 | Wrong population |
| No population-level outcomes | 394 | Wrong outcome |
| Climate not primary driver | 312 | Wrong exposure |
| Theoretical/modeling only | 284 | Wrong study design |
| Narrative reviews/opinion pieces | 267 | Wrong study type |
| Mean warming only/not variability or extremes | 467 | Other |
| Total excluded | 2,210 | |
| Full texts assessed | 460 | |
| ✔ 2,670−2,210 = 460 (matches PRISMA) | ||
| Table S4C: Full-text exclusions with reasons (n = 320). | ||
| Reason for Exclusion | Number | PRISMA Category |
| Mean climate trends only | 72 | Wrong exposure |
| No empirical population metrics | 68 | Wrong outcome |
| Grey / non-peer-reviewed sources | 61 | Wrong study type |
| Plant-only or inseparable mixed systems | 47 | Wrong population |
| Insufficient methodological detail | 38 | Other |
| Duplicate dataset | 34 | Other |
| Total excluded | 320 | |
| ✔ 460−320 = 140 (matches PRISMA) | ||
This table lists all peer-reviewed empirical studies contributing to the qualitative and quantitative synthesis. Only studies restricted to animal taxa and explicitly addressing climate variability and/or climatic extremes were included.
| Table S4D: Studies included in the systematic review (n = 140). | ||||||
| Study | Taxon | Realm | Dominant Climate Driver | Population Metric(s) | Study Design | Quality Score |
| Visser et al.19 | Birds | Terrestrial | Temperature variability | Breeding phenology, fitness | Longitudinal empirical | High |
| Paniw et al.6 | Mammals | Terrestrial | Weather variability | Survival, reproduction | Multi-species demographic | High |
| Franks et al.21 | Birds | Terrestrial | Temperature variability | Survival, reproduction | Long-term field study | High |
| Flores et al.20 | Mammals | Terrestrial | Extreme heatwaves | Reproductive timing | Longitudinal empirical | High |
| Lenzi et al.22 | Amphibians | Freshwater / terrestrial | Temp & precipitation variability | Recruitment | Long-term monitoring | High |
| Shimizu et al.18 | Insects | Terrestrial (alpine) | Temperature variability | Population growth | Field + monitoring | High |
| Thackeray et al.41 | Multiple animal taxa | Terrestrial / freshwater | Seasonal variability | Phenology | Meta-analysis | High |
| Block et al.26 | Marine vertebrates | Marine | Ocean variability | Migration | Telemetry-based | High |
| Singh et al.29 | Large mammals | Terrestrial | Temperature variability | Seasonal movements | GPS tracking | High |
| Parmesan and Yohe40 | Multiple taxa | Global | Climate variability | Range shifts | Comparative empirical | High |
| Leigh et al.10 | Vertebrates & invertebrates | Global | Climate extremes | Genetic diversity | Meta-analysis | High |
| Willi et al.11 | Multiple taxa | Terrestrial | Climatic extremes | Adaptive potential | Experimental + synthesis | High |
| López-Gatius et al.35 | Mammals | Terrestrial | Temperature variability | Genetic diversity | Empirical + modeling | Medium |
| … | … | … | … | … | … | … |
Only peer-reviewed empirical or data-driven synthesis studies were included.
- Grey literature, news media, and non-peer-reviewed preprints were excluded at full-text screening.
- Each study represents one analytical unit in proportional summaries.
- Quality scores informed interpretive weighting only (Supplementary Table S3).








