From a hefty greenery to a parched paradise: assessing the impacts of climate change on water security and biodiversity decline in the Western Cape Province of South Africa

Background: This systematic review integrates existing evidence within a multi-scale analytical framework to understand how climate-induced water stress affects biodiversity in Mediterranean-climate regions, using Erica species in the Cape Floristic Region (CFR) as a model system.

Methods: Following PRISMA guidelines, we systematically reviewed 57 peer-reviewed articles from Scopus and Google Scholar, organizing findings using the DPSIR (Drivers-Pressures-State-Impacts-Responses) framework across three spatial scales: global Mediterranean, regional Sub-Saharan Africa, and local Cape Floristic Region.

Results and Discussion: Systematic synthesis reveals water availability as the primary mechanism mediating climate impacts on Erica biodiversity across all examined contexts. Integration of quantitative evidence from multiple independent studies identifies threshold patterns where water deficits of 20%–30% relative to historical conditions distinguish resilient from vulnerable populations in European systems, though documented thresholds vary among populations (20%–40% range) and require validation for CFR endemic species. Cascading impacts progress from physiological stress (40% flowering reduction under experimental drought) through demographic bottlenecks (50%–70% germination decline under moisture limitation) to ecosystem functional changes. While physiological response mechanisms operate consistently across Mediterranean regions, vulnerability magnitude is context-dependent: synthesis suggests CFR’s approximately 700 endemic species exhibit narrower tolerances than European congeners, reflecting evolution under stable climatic conditions versus historical variability. This multi-scale framework distinguishes generalizable physiological principles from context-specific vulnerabilities, providing operational guidance for conservation priority-setting. The DPSIR structure explicitly traces causal pathways from global drivers to local responses, enabling identification of intervention leverage points across organizational levels from regional water policy through landscape connectivity to site-scale microhabitat management. Findings indicate that conservation strategies developed for European Erica populations may underestimate CFR vulnerabilities without accounting for narrower endemic tolerances and limited adaptive capacity arising from rapid recent diversification.

1 Introduction

Climate change is one of the most important and urgent environmental issues, which drastically alters the patterns of global temperature and precipitation (Bellard et al., 2012; Habibullah et al., 2022). According to recent projections by the Intergovernmental Panel on Climate Change, global temperatures may increase by up to 6.4 °C by the end of this century under sustained greenhouse gas emission scenarios (Reinman, 2012). These profound climatic shifts significantly impact hydrological cycles, water resource availability, and ecosystem functionality across diverse geographical and ecological contexts (Bhaga et al., 2020; du Plessis and du Plessis, 2017).

The availability of water resources is a crucial intersection of biodiversity responses and climate change, often forming the main barrier to plant productivity and ecosystem vitality (Gan et al., 2021; Lambers et al., 2008). Climate-driven changes in thermal and hydrological regimes directly control plant physiological function, phenology, and species range dynamics. These changes also affect the volume and timing of water (Gao et al., 2016; Zhang et al., 2015). Mediterranean regions are especially vulnerable due to their regular seasonal drought. With projected aridification, these areas face greater risks to plant communities and vital ecosystem services (Chen et al., 2011; Chen et al., 2020). Aridity is a central driver across Mediterranean biomes. For example, in North Africa, aridity gradients structure the floristic diversity linked to Erica arborea (Djillali et al., 2023). This confirms water availability as a key pressure shaping Erica’s distribution in many Mediterranean settings. Sub-Saharan Africa faces unique challenges, with expected temperature increases of 1.1 °C–6.4 °C by 2100, compared to 1980–1999 levels (Reinman, 2012). Rising temperatures, shifting rainfall, and more frequent extremes produce many vulnerabilities in African water resources and biodiversity (Thornton et al., 2014; Serdeczny et al., 2017). Diverse biomes, ranging from East African Afromontane systems to Southern African fynbos zones, face unique climate stressor combinations that may surpass the adaptive potential of indigenous flora and fauna (Hannah et al., 2014). Therefore, a thorough understanding of these regionally specific climate-water-biodiversity relationships is necessary for effective conservation planning (Midgley et al., 2002; Midgley et al., 2003).

Despite growing recognition of climate change threats to biodiversity, existing review literature exhibits important limitations in addressing water-mediated climate impacts across spatial scales. At the global scale, recent reviews have examined climate-biodiversity relationships broadly (Bellard et al., 2012; Pecl et al., 2017) and Mediterranean ecosystem responses specifically (Peñuelas et al., 2017), yet these syntheses have not systematically identified water availability as a primary mediating mechanism linking climate drivers to biodiversity impacts across organizational levels. Bellard et al. (2012) synthesis documented widespread climate impacts on species distributions and phenology across taxonomic groups and biomes, establishing climate change as a major biodiversity threat. However, their review did not elucidate mechanistic pathways through specific environmental alterations, such as hydrological changes, which limit guidance for targeted conservation interventions. Similarly, Pecl et al. (2017) synthesized evidence for climate-driven species redistribution globally yet focused on distributional shifts rather than the physiological and demographic mechanisms underlying range changes in water-limited systems.

Peñuelas et al. (2017) comprehensive review of Mediterranean forest responses to global change identified drought intensification, altered fire regimes, and increased pest pressure as key stressors affecting ecosystem functioning and biodiversity. While valuable for establishing regional context and recognizing drought as an important pressure, their synthesis did not integrate evidence across spatial scales to distinguish which patterns represent generalizable Mediterranean responses versus vulnerabilities specific to particular evolutionary or climatic contexts. This gap is particularly important given that Mediterranean-climate regions exhibit substantial differences in evolutionary history, with implications for adaptive capacity: the Cape Floristic Region’s flora diversified rapidly over the last 10,000 years under stable climatic conditions (Cowling et al., 2009), whereas Mediterranean Basin species evolved under greater Quaternary climate variability (Médail and Diadema, 2009).

Hannah et al. (2014) introduced microrefugia, stepping-stones, and holdouts as frameworks for understanding fine-scale species responses to climate change. Their analysis provided valuable theoretical tools for conservation planning, particularly emphasizing the role of topographic and microclimatic heterogeneity in facilitating persistence. However, their framework lacks integration with water resource dynamics and does not provide operational guidance for identifying and managing microrefugia in water-limited Mediterranean systems where soil moisture rather than temperature may determine microsite suitability. For plant communities specifically, existing reviews have not systematically examined how water availability mediates climate impacts across the complete hierarchy from individual physiological responses through population demography to ecosystem functioning.

Early syntheses of climate adaptation strategies provided important foundations but revealed persistent implementation gaps. Heller and Zavaleta (2009) systematically reviewed 22 years of climate-biodiversity adaptation recommendations across 113 papers, revealing that while landscape connectivity and regional coordination were frequently advocated, 70% of recommendations remained general principles lacking operational specificity. Critically, their synthesis identified that while water availability was “frequently mentioned” as a climate change pressure, existing reviews had not systematically identified it as a primary mediating mechanism or traced its effects across organizational levels from physiology through demography to ecosystem functioning. This mechanistic gap limits understanding of where and how interventions might most effectively buffer climate impacts.

Building on this foundation, Mawdsley et al. (2009) provided a comprehensive taxonomy of 16 adaptation strategies organized into four functional categories (land/water protection, species management, monitoring/planning, and law/policy), with a critical evaluation of each approach’s strengths and limitations. Their seminal observation that conservation practitioners possess many necessary tools but must apply them “in novel and innovative ways” to address climate change highlighted the implementation challenge. However, while their functional categorization effectively organized adaptation tools, it did not systematically address how these strategies operate across spatial scales or trace mechanistic pathways linking climate drivers to biodiversity responses through specific environmental mediators. More recently, Xu et al. (2025) advanced cross-scale frameworks by demonstrating how adaptive management strategies at regional, landscape, and site scales interact through both ecological cascades and policy implementation mechanisms, providing theoretical validation for multi-level conservation coordination.

Fagúndez (2013) examined heathland responses to global change in European systems, noting threats from climate change, nitrogen deposition, and altered fire regimes specifically for the Erica genus. Pirie et al. (2022) synthesis outlined conservation priorities for Erica globally, emphasizing ex situ conservation through seed banking and living collections, given extinction risks. More recently, Pirie et al. (2025) conducted a systematic conservation gap analysis for Erica, providing essential evidence to re-prioritize resource allocation to geographically and taxonomically underserved populations. While these syntheses provide important genus-specific context, they have not systematically integrated evidence across the complete spatial hierarchy from global Mediterranean patterns through sub-Saharan African regional dynamics to Cape Floristic Region endemic vulnerabilities. Taken together, these reviews establish important foundations but reveal persistent gaps when attempting to trace water-mediated climate impacts across both spatial scales and organizational levels.

Four specific gaps persist in existing syntheses despite these advances. First, while water availability is frequently mentioned as a climate change pressure, existing reviews have not systematically identified it as the primary mediating mechanism and traced its effects across organizational levels from physiology through demography to ecosystem functioning. This mechanistic gap limits understanding of where and how interventions might most effectively buffer climate impacts. Second, existing syntheses have not applied explicit analytical frameworks (such as DPSIR) to systematically organize causal pathways from global drivers through regional pressures to local responses across multiple scales, limiting their utility for multi-level conservation planning that must coordinate interventions across governance scales. Third, and most critically for endemic-rich systems, previous reviews have not explicitly distinguished generalizable physiological mechanisms operating consistently across Mediterranean regions from context-specific vulnerabilities arising from regional evolutionary histories and endemism patterns. This final gap is particularly important given that global climate-biodiversity models calibrated primarily on temperate systems with broad species ranges systematically underestimate risks for recently diversified endemic-rich floras with narrow tolerances (Yates et al., 2010). Fourth, despite the recognition that conservation requires “novel application” of existing tools (Mawdsley et al., 2009) and coordination across scales (Xu et al., 2025), operational guidance for translating these principles into context-specific, mechanistically informed conservation strategies remains limited, particularly for identifying intervention leverage points across the driver-pressure-state-impact cascade.

These gaps are particularly consequential for the Cape Floristic Region (CFR), one of the world’s six floral kingdoms and a global biodiversity hotspot. Hosting over 9,000 vascular plant species with 69% endemism (Mittermeier et al., 2011; Manning and Goldblatt, 2012), the CFR’s exceptional plant diversity, including approximately 700 endemic Erica species, provides both conservation urgency and research opportunity for investigating climate change impacts on taxonomically diverse plant communities (Cowling et al., 2009; Pirie et al., 2016). Beyond their ecological role in fire-prone ecosystems, the Erica genus holds significant socio-economic value. Comprehensive reviews on Mediterranean species confirm their extensive ethnomedicinal use and rich phytochemical composition, highlighting potent anti-inflammatory and anti-Alzheimer’s potential (Jabal et al., 2025). This biological value highlights the extensive effects of biodiversity loss due to climate change on humans. Rising temperatures, altered precipitation patterns, and an increase in extreme weather events heighten South Africa’s vulnerability to water resources and biodiversity. Plant genera with expansive geographic ranges yet significant regional diversification, such as Erica, offer exceptionally useful model systems for addressing the identified gaps in multi-scale climate change research (Ojeda et al., 2016). With species occupying diverse habitats across Europe, Africa, and Mediterranean-climate regions globally, Erica taxa display varied physiological adaptations, ecological functions, and conservation statuses (Pirie et al., 2016; Pirie et al., 2017). These characteristics facilitate comparative analyses of climate change responses across geographical contexts, from global distribution patterns to regional community dynamics and local population viability (Bellard et al., 2012).

Specifically, Erica species provide an ideal model system for addressing the identified gaps because: (1) their global distribution across Mediterranean-climate regions enables assessment of generalizable physiological mechanisms and thresholds reported in existing literature; (2) pronounced regional diversification, with approximately 700 endemic CFR species versus widespread European congeners permits direct comparison of context-specific vulnerabilities related to evolutionary history; and (3) documented sensitivity to water availability across multiple independent studies provides empirical basis for tracing mechanistic pathways across organizational levels from individual stress responses through population dynamics to ecosystem functioning. Moreover, Erica species’ functional roles as cornerstone taxa in fynbos systems (Cowling et al., 2015; Hitchcock and Rebelo, 2017) mean that understanding their responses to water stress has direct implications for broader ecosystem resilience and conservation planning. The challenge of distinguishing generalizable mechanisms from context-specific vulnerabilities is exemplified by species like Erica sicula, whose distribution is broad but highly disjunct across the Mediterranean (Pasta et al., 2025), complicating traditional risk assessments and mandating coordinated global conservation strategies for geographically isolated, locally endangered populations.

The overarching objective of this systematic review is to integrate existing evidence within a multi-scale analytical framework that identifies both generalizable ecological principles of plant responses to climate-induced water stress and context-specific vulnerabilities associated with regional evolutionary and climatic distinctiveness, ultimately informing evidence-based conservation prioritization for water-dependent plant communities in Mediterranean-climate biodiversity hotspots. This objective is implemented through four specific interrelated aims. First, to synthesize evidence on universally applicable patterns of climate-water-biodiversity interactions across global Mediterranean-climate regions, identifying common physiological thresholds and response mechanisms reported for Erica species. Second, to integrate findings regarding scale-dependent differences in species responses to water availability changes, examining how regional climate dynamics (sub-Saharan Africa) and local evolutionary contexts (Cape Floristic Region) modify general patterns. Third, to trace from existing literature the mechanistic pathways by which climate change propagates from global atmospheric forcing through regional hydrological alterations to local biodiversity effects, including cross-scale transmission processes. Fourth, to assess conservation implications by integrating evidence on physiological thresholds, adaptive responses, and ecosystem functioning to generate priority-setting criteria for conserving threatened Erica communities under future climatic scenarios.

This review employs three complementary analytical techniques that extend insights from current frameworks to address the identified gaps, building directly upon the foundations established by Poiani et al. (2000), Heller and Zavaleta (2009), Mawdsley et al. (2009), and Xu et al. (2025). First, we systematically trace water availability’s role as a mediating mechanism across organizational levels from individual physiological responses through population demography to ecosystem functioning, providing mechanistic specificity that broader syntheses have not achieved (Bellard et al., 2012; Pecl et al., 2017). This addresses Heller and Zavaleta (2009) finding that water was “frequently mentioned” but not systematically analysed as a primary mediating pathway, providing focused analytical effort on synthesizing evidence regarding the most critical limiting factor for Mediterranean plant communities and documenting how its effects propagate through biological hierarchies.

Second, we apply the DPSIR (Drivers-Pressures-State-Impacts-Responses) framework to explicitly organize causal pathways from global atmospheric forcing through regional hydrological alterations to local biodiversity responses across multiple spatial scales. Following Heller and Zavaleta (2009) and Mawdsley et al. (2009) identification of the need for practical adaptation planning processes that integrate recommendations across scales, this structured approach extends typical single-scale applications of DPSIR to provide a systematic organization for multi-level conservation planning, directly addressing the second gap by creating transparent linkages between climate drivers and management responses across governance scales. While Poiani et al. (2000) provided an influential framework for designing functional conservation areas based on maintaining ecosystems and species within their natural ranges of variability across multiple spatial scales, their framework requires region-specific and taxon-specific empirical evidence. This synthesis addresses these evidence gaps for water-dependent Mediterranean plant communities, providing the mechanistic understanding and quantitative thresholds necessary to operationalize multi-scale conservation frameworks like Poiani et al. (2000) in the context of accelerating climate change.

Third, we explicitly distinguish generalizable physiological principles operating consistently across Mediterranean regions from context-specific vulnerabilities arising from the Cape Floristic Region’s unique evolutionary history, specifically, rapid recent diversification under stable climates producing approximately 700 endemic Erica species with potentially narrow tolerances. By facilitating the identification of both site-specific elements necessitating localized conservation strategies and universal patterns applicable across Mediterranean systems, this scale-dependent analytical structure fills the third gap. This multi-scale, mechanistically grounded approach operationalizes Mawdsley et al. (2009) call for “novel and innovative” application of conservation tools by demonstrating how established strategies (riparian restoration, corridor protection, species translocation) require context-specific parameterization based on water-mediated thresholds and evolutionary history, while extending Xu et al. (2025) theoretical cross-scale framework into practical conservation guidance.

This paper investigates how climate-induced shifts in water availability affect biodiversity across multiple scales, using the widespread yet ecologically significant Erica genus as a model system for understanding responses from physiological stress to ecosystem-level impacts (West et al., 2012). Through systematic synthesis of 57 peer-reviewed studies organized within the DPSIR framework, we demonstrate that while physiological mechanisms reported across studies are generalizable, vulnerability magnitude is context-dependent, with direct implications for prioritizing conservation investments in Mediterranean-climate biodiversity hotspots facing accelerating climate change.

2 Materials and methods

In accordance with PRISMA guidelines, we carried out a systematic literature review to compile data on climate change effects on biodiversity conservation and water resources (Moher et al., 2009). In June and July 2025, searches were conducted using Google Scholar and Scopus databases. We used search terms: (climate change OR global warming) AND (water OR precipitation OR drought) AND (biodiversity OR Erica OR plant communities) AND (Mediterranean OR “Cape Floristic Region” OR “Sub-Saharan Africa”). The combined search yielded 490 articles (n = 238 Scopus, n = 252 Google Scholar). After removing duplicates (n = 67), 423 articles remained for screening. A three-stage screening process was implemented following Liberati et al. (2009). Title screening excluded 257 irrelevant articles. Abstract screening of the remaining 166 articles against the inclusion criteria (Table 1) identified 49 articles meeting all criteria. Following Pullin and Stewart (2006) guidelines, we examined reference lists, identifying 17 additional papers. Screening identified eight meeting criteria, increasing the final sample to 57 articles (Figure 1).

Table 1

Table 1. Inclusion and exclusion criteria for systematic review.

Figure 1

Figure 1. PRISMA flow diagram illustrating the systematic selection process from the initial database searches through final inclusion.

2.1 Analytical framework

The synthesis employed two complementary frameworks. First, DPSIR (Kristensen, 2004) structured analysis tracing causal pathways: Drivers (greenhouse gas emissions, land use change), Pressures (temperature increase, altered precipitation), State (water availability, vegetation health), Impacts (species loss, phenological shifts), and Responses (conservation strategies). DPSIR was selected for explicitly organizing causal chains, providing a systematic structure for multi-level planning, addressing Heller and Zavaleta (2009) call for practical frameworks distinguishing impact assessment from implementation. Second, a multi-scale spatial framework was employed. Building upon landscape ecology principles (Xu et al., 2025; Poiani et al., 2000), recognizing that climate impacts operate across nested scales requiring coordinated responses, studies were categorized as: (1) Global scale (Mediterranean-climate regions worldwide); (2) Regional scale (Sub-Saharan African climate dynamics); (3) Local scale (CFR endemic Erica species). Our geographic-scale framework differs from Poiani et al. (2000) ecological-scale framework. However, both recognize that multi-scale understanding is essential. Our synthesis informs ecological-scale planning that Poiani et al. (2000) framework guides identified water thresholds provide an empirical basis for determining minimum dynamic area requirements for fire-adapted fynbos. Articles were assigned to spatial scales based on primary geographic focus and finest spatial resolution. Scale assignments were reviewed by both authors, with disagreements resolved through discussion.

2.2 Thematic synthesis

Thematic synthesis, involving an inductive and iterative coding technique (Thomas et al., 2012), served as the primary analytical approach to systematically identify, categorize, and synthesize key findings across the included literature. The first author performed the initial coding. To minimize the subjectivity inherent to single-coder analysis, we implemented multiple quality assurance processes. A structured codebook was initially generated via the detailed, line-by-line analysis of five articles, which were purposively selected to represent the full spectrum of geographical scales (global/regional/local) and study types (experimental/observational/modeling). This initial coding resulted in a preliminary codebook with working definitions for all codes. Second, the codebook was refined through three iterations across the complete dataset. In the first iteration (n = 20 articles), 47 initial codes were identified. The second iteration (n = 20 articles) involved aggregation of overlapping codes (e.g., “reduced flowering” and “delayed flowering” combined into “phenological impacts”) and differentiation of overly broad codes (e.g., “water stress” decomposed into “drought duration,” “seasonal water deficit,” and “soil moisture depletion”), resulting in 38 refined codes. The third iteration (n = 17 articles) finalized 35 codes organized under seven thematic clusters corresponding with DPSIR framework components and research objectives.

Third, at each stage of the iterative process, the second author (supervisor with expertise in climate change ecology and systematic reviews) reviewed the coding scheme and emerging themes, enhancing inter-rater reliability and reducing individual interpretation bias. After the coding framework was stabilized, individual codes were aggregated into higher thematic categories through the interactive process of abstraction (Thomas et al., 2012). For instance, codes including “germination failure,” “seedling mortality,” and “recruitment bottlenecks” were synthesized into the overarching theme “life cycle vulnerabilities.” The DPSIR framework systematically organizes causal pathways from anthropogenic drivers to conservation responses (Figure 2). This multi-scale structure enables transparent linkages between global climate forcing and local biodiversity impacts across nested spatial hierarchies. Through this process, seven key themes emerged that map directly to the research objectives and DPSIR framework: (1) Global climate-water-biodiversity patterns; (2) Sub-Saharan African regional dynamics; (3) Cape Floristic Region-specific vulnerabilities; (4) Physiological response mechanisms; (5) Population and community-level impacts; (6) Ecosystem process and function changes; and (7) Conservation and management responses. The resulting synthesis is organized around these themes to systematically address each research objective.

Figure 2

Figure 2. DPSIR framework illustrating nested interactions between global climate drivers, regional pressures, local ecosystem states, species-level impacts, and cross-scale conservation responses.

2.3 Limitations

A number of methodological constraints should be taken into consideration when interpreting review results. First, relying on a single coder increases the risk of individual interpretive bias even when quality assurance measures like explicit codebook documentation and supervisor validation are used. However, the transparency of the coding process and detailed codebook documentation enable independent evaluation and replication. Second, as this is a qualitative synthesis, a quantitative meta-analysis was not performed. Although this approach was necessary to accommodate the breadth of data types, study designs, and spatial scales examined, it necessarily restricts the statistical rigor and strength of causal claims that can be made regarding climate-species relationships. Third, the restriction to English-language peer-reviewed literature may introduce language and publication bias, potentially excluding relevant findings published in other languages or formats such as technical reports and grey literature. Fourth, the search strategy, while comprehensive within the selected databases, was limited to Scopus and Google Scholar and did not include all potential sources (e.g., Web of Science, discipline-specific databases), which may have resulted in the omission of relevant studies. Fifth, the geographic and temporal distribution of included studies reflects the existing research landscape, which may be biased toward certain regions (particularly European Mediterranean systems, where Erica research has a longer history) and recent time periods, potentially affecting the generalizability of synthesized patterns across all Mediterranean-climate systems and evolutionary contexts. Lastly, only 21% reviewed studies reported formal statistical relationships between species responses and climate variables, which limits our capacity to synthesize effect sizes quantitatively and necessitates relying on qualitative synthesis of reported patterns and trends.

3 Results

3.1 Generalizable patterns across global Mediterranean-climate regions

Climate change significantly influences global biodiversity through impacts on species’ life cycles, physical characteristics, and geographical distributions (Muluneh, 2021; Habibullah et al., 2022). Studies demonstrate how these changes reshape biological communities, accelerating biodiversity loss worldwide (Bellard et al., 2012; Pecl et al., 2017). The genus Erica constitutes a keystone component of ecosystem architecture, demonstrating particular prominence across Mediterranean zones (Fagúndez, 2013). Beyond ecological roles, Erica holds significant socio-economic value through ethnomedicinal use and phytochemical composition (Jabal et al., 2025; Adu-Amankwaah et al., 2025). These taxa provide essential ecosystem services through extensive belowground structures facilitating hydrological regulation (Brempong et al., 2023). Pre-eminent global temperatures have restructured precipitation regimes, with Mediterranean-type ecosystems demonstrating pronounced susceptibility (Peñuelas et al., 2017). Studies have documented substantial deviations in rainfall attributes directly controlling water resource availability (Chen et al., 2020; Gao et al., 2016). These perturbations manifest as extended aridity periods, diminished soil moisture retention, and amplified evapotranspiration rates (Chen et al., 2020; Gao et al., 2016). The primary mechanism of climate impact is mediated by water availability. Convergent evidence demonstrates that drought directly constrains the structural diversity of wood in southern African Erica (Akinlabi et al., 2023).

Multiple independent studies consistently document Erica species exhibit marked sensitivity to thermal and precipitation gradients modulating phenological dynamics and reproductive fitness (Prieto et al., 2008; Prieto et al., 2009; Bernal et al., 2011). Prieto et al. (2008) revealed pronounced dependence of flowering phenology on precipitation regimes, with experimental manipulations simulating moisture limitation consistently yielding temporal displacement and quantitative reduction in flowering.

Water resource availability represents a fundamental determinant of plant development and ecosystem operation (Lambers et al., 2008; Gan et al., 2021). The analytical framework indicates climate-mediated water stress negatively influences biodiversity through multiple interacting pathways (Chen et al., 2020; Gao et al., 2016). Erica populations consistently demonstrate robust associations between precipitation regimes and reproductive success (Prieto et al., 2008). Complementary studies by Nogués et al. (2012) and Bernal et al. (2011) indicated physiological response to combined drought and thermal stressors exhibits seasonal differences, with water deficit effects exacerbated during summer months corresponding to maximum evaporative demand. Geographic heterogeneity in climate sensitivity was documented. Chamorro et al. (2018) determined climate-induced thermal increases disproportionately affect higher-latitude populations where elevated temperatures inhibit seed germination. Conversely, Llorens et al. (2004) noted heightened susceptibility to increased temperature and moisture deficiencies in northern populations relative to southern populations. Global literature consistently indicates climate change restructures ecological communities via multifaceted mechanisms transcending single-species responses. Progressive modification of biotic interactions is driven by climate-induced fluctuations in water availability (Folke et al., 2004; Ortega et al., 2024). In North African systems, floristic diversity associated with Erica arborea is fundamentally structured by aridity gradients (Djillali et al., 2023; Boughediri et al., 2025), confirming water availability as central pressure shaping Erica distribution across Mediterranean biomes.

Integration of quantitative relationships (Table 2) identifies three consistent patterns. First, precipitation demonstrates consistently positive effects on Erica growth and reproduction across all contexts (Spain: Prieto et al., 2008; Ethiopia; Jacob et al., 2020; South Africa: Brown et al., 2016), establishing water availability as the primary constraint. Second, temperature effects are context-dependent and conditional on water availability: positive in moist conditions (Jacob et al., 2020), negative under stress (Nogués et al., 2012). Third, climate sensitivity varies between populations (Llorens et al., 2004; Chamorro et al., 2018). Although formal correlation coefficients are rarely reported (only 17%), consistent directional responses provide substantial qualitative evidence for water availability as the primary mechanism. Convergent evidence suggests water deficits of 20%–30% relative to historical conditions may represent critical thresholds distinguishing resilient from vulnerable populations, though constrained by: (1) derivation primarily from two European studies (Prieto et al., 2008; Llorens et al., 2004), (2) documented variation 20%–40% across populations, and (3) limited testing across CFR endemics. This synthesis-derived threshold should be interpreted as a working hypothesis requiring validation rather than a universally applicable parameter.

Table 2

Table 2. Quantitative synthesis of Erica species responses to climate variables.

The convergent evidence suggesting water deficits of 20%–30% provides an initial estimate of the natural range of variability (Poiani et al., 2000) for precipitation regimes supporting Erica populations. Poiani et al. (2009) defined functional conservation areas as maintaining “focal ecosystems within natural ranges of variability” over 100–500-year periods. The documented threshold where populations transition from resilient to vulnerable represents the boundary of natural variability that CFR species can tolerate. Maintaining precipitation within this historical envelope should constitute the primary conservation objective, as exceedance would push populations beyond the adaptive capacity evolved under stable Holocene conditions. However, these thresholds derive primarily from European studies examining species evolved under greater Quaternary variability. Whether CFR endemic species exhibit similar or narrower tolerance ranges remains an empirical question requiring direct experimental validation.

3.2 Regional and local vulnerabilities

In Sub-Saharan Africa, climate change manifests through temperature increases of 0.7 °C during the 20th century (Ring et al., 2012; Marcott et al., 2013), with projections of 1.1 °C–6.4 °C by 2100 (Intergovernmental Panel on Climate Change (IPCC), 2014). Regional vulnerability includes significant rainfall pattern variations and increases extreme weather frequency (Thornton et al., 2014; Serdeczny et al., 2017). Jacob et al. (2020) dendrochronological analysis of E. arborea in Ethiopian highlands revealed that temperature effects were modulated by water availability: tree growth increased during years with adequate rainfall but declined during drought years. This conditional response indicates warming influences precipitation effect magnitude. Scenarios with maintained rainfall may allow Erica persistence, whereas warming coupled with precipitation declines will cause threshold exceedance. Based on the synthesis of experimental evidence (Prieto et al., 2008; Llorens et al., 2004), a tentative critical threshold occurs when evapotranspiration demand exceeds water supply by 20%–30%. However, this requires validation across Sub-Saharan African populations.

In East Africa, Erica species are critical components of upper treeline ecotones, with significant populations in Ethiopian highlands (Jacob et al., 2020). Research established a complex relationship between growth dynamics and hydrological regimes, showing a significant positive correlation between growth and precipitation during the primary growing season. Elevated temperatures promote growth contingent upon sufficient moisture; conversely, higher temperatures exert an inhibitory effect during drought. Literature reveals that distribution and survival face complex challenges attributed to climate change. Kidane et al. (2022) projected climate change would lead to upward distributional shifts and community composition alterations, indicating potential altitudinal shifts, profound composition changes, habitat fragmentation, and critical moisture availability influence (Vicente-Serrano et al., 2013).

The Cape Floristic Region hosts over 9,000 vascular plant species with 69% endemism (Mittermeier et al., 2011; Manning and Goldblatt, 2012). The CFR’s high susceptibility stems from exceptional biodiversity evolved under specialized historical climatic conditions (Adedoja, 2019). The genus Erica, harbouring approximately 700 endemic species (Pirie et al., 2023), functions as a cornerstone taxon (Cowling et al., 2015) and a keystone species (Hitchcock and Rebelo, 2017), playing critical roles in water regulation, carbon sequestration, and soil stabilization. Pirie et al. (2024) demonstrated significant spatial decoupling of species richness and phylogenetic diversity, confirming conservation planning must explicitly incorporate phylogenetic diversity. Historical data reveal significant warming trends, with Kruger and Shongwe (2004) documenting temperature increases 1960–2003, Warburton et al. (2005) identifying faster warming 1980–2000 versus 1950–1970, and Haensler et al. (2010) observing 0.1 °C–0.2 °C per decade increases 1901–2006. Multiple bioclimatic models indicate even minor temperature increases significantly affect plant growth and reproductive success (Midgley et al., 2002, 2003; Bomhard et al., 2005). Dawson et al. (2012) demonstrated a direct correlation between drought and reduced flowering intensity. Brown et al. (2016) showed that low moisture reduces germination rates, establishing recruitment barriers. Water stress also detrimentally affects post-germination seedling development. Road fragmentation interferes with important mutualisms like bird pollination in Erica glandulosa (Grobler and Campbell, 2022). West et al. (2012) project that anticipated precipitation alterations will force numerous Erica species to surpass physiological thresholds. Manzano et al. (2023) demonstrated cascading consequences compromising survival and reproductive success. Wilson et al. (2015) emphasized Erica species evolved within narrow thermal ranges, with temperature increases pushing taxa beyond physiological tolerances.

3.3 Mechanistic pathways linking climate drivers to biodiversity impacts

Erica taxa display varying susceptibility to climate stressors, with consistent evidence demonstrating that drought diminishes reproductive output through reduced flowering (Dawson et al., 2012; Brown et al., 2016). Phenological shifts risk disrupting ecological interactions with pollinators and seed dispersers. Low moisture availability results in severe germination reduction and impedes seedling establishment, suggesting climate change simultaneously impacts multiple developmental stages. Projected precipitation alterations (West et al., 2012) will intensify challenges by fundamentally changing water availability patterns, threatening to push species beyond physiological limits. Water stress exhibits cascading consequences extending beyond individual survival to impair reproductive success, population dynamics, and distribution patterns (Manzano et al., 2023; Yates et al., 2010). Temperature modifications present substantial threats as many Erica species evolved within narrow thermal ranges (Wilson et al., 2015; Pirie et al., 2022). Findings across spatial scales can be systematically organized within the DPSIR framework to clarify causal pathways from anthropogenic forcing to conservation action (Table 3).

Table 3

Table 3. DPSIR framework synthesis of climate-water-biodiversity relationships for Erica communities.

3.4 Conservation implications from integrated evidence

Synthesis across scales reveals critical priorities. At the global Mediterranean scale, a consistently documented positive precipitation-reproduction relationship (Table 2) establishes water availability as the primary conservation target. Management should prioritize maintaining adequate moisture during critical phenological windows, particularly flowering periods, when water deficits produce 40% reproductive output reductions (Prieto et al., 2008). Identification of seasonal vulnerability patterns, with summer drought producing disproportionate impacts (Nogués et al., 2012; Bernal et al., 2011), indicates conservation resources should be strategically allocated to mitigate water stress during peak evaporative demand periods. Documented population-level variation in climate sensitivity (Llorens et al., 2004; Chamorro et al., 2018) necessitates genetically informed conservation. Northern European populations exhibiting heightened sensitivity may serve as early warning systems, while southern populations demonstrating resilience could provide source material for assisted gene flow or ex situ programs. At the sub-Saharan African scale, conditional temperature effects (Jacob et al., 2020) reveal conservation success depends critically on integrated water resource management. The projected 1.1 °C–6.4 °C warming (Intergovernmental Panel on Climate Change (IPCC), 2014) will exceed physiological thresholds only if coupled with precipitation declines.

Topographic complexity offers both vulnerability and opportunity for East African Afromontane populations. While climate-driven upslope shifts (Kidane et al., 2022) may lead to mountaintop extinctions, topographic heterogeneity creates microrefugia where local moisture availability buffers regional trends. Conservation planning should map microrefugia through fine-scale hydrological modeling, prioritizing moisture-retaining landscape positions (north-facing slopes, valley bottoms). For CFR, the confluence of exceptional endemism, rapid historical warming (Haensler et al., 2010), and narrow physiological tolerance creates an acute conservation imperative. Documented impacts across multiple life stages (germination failure, seedling mortality, recruitment bottlenecks, reduced flowering: Dawson et al., 2012; Brown et al., 2016) indicate population viability hinges upon successful demographic transitions. Conservation interventions must target stage-specific vulnerabilities, including water supplementation during germination, microsite management for establishment, and pollinator protection to maximize seed set during flowering stress.

Beyond species preservation, maintaining Erica community function is essential for ecosystem resilience. As cornerstone taxa (Cowling et al., 2015; Hitchcock and Rebelo, 2017), these plants deliver critical services (water regulation, carbon sequestration, soil stabilization); their loss would precipitate cascading impacts through soil chemistry and microhabitat disruption. Documented functional diversity within Erica communities (West et al., 2012) represents a vital insurance mechanism against climate uncertainty. Therefore, functional diversity preservation should take precedence over species richness. Translating the DPSIR framework into actionable responses requires a targeted, data-driven approach addressing specific shortfalls identified by systematic conservation gap analysis (Pirie et al., 2025). Given documented pharmacological importance (Jabal et al., 2025; Adu-Amankwaah et al., 2025), targeted ex-situ conservation efforts and chemical screening should be prioritized.

Adoption of adaptive management frameworks is essential given the conditional and nonlinear nature of climate-water-biodiversity relationships. These frameworks should incorporate threshold-based triggers, ongoing monitoring, and adaptable intervention techniques. This adaptive approach is particularly critical given that only 17% of reviewed studies report formal statistical relationships (Table 2), and proposed thresholds represent synthesis-derived hypotheses requiring validation. To assist decision-makers in identifying the approach of the 20%–30% deficit threshold in real time, a monitoring timetable should be established. This timetable aligns ecological indicators such as soil moisture levels, phenological changes, and species stress signals with policy action windows, ensuring timely interventions. Water availability has emerged as the primary mediator; therefore, biodiversity conservation must integrate holistically with water resource management. Protected area management must adopt hydrological considerations, including natural flow regime maintenance, groundwater abstraction restriction, and fire management, ensuring water-retaining vegetation preservation.

4 Discussion

To clarify climate change impacts on water-dependent plant communities, this systematic review combined data from 57 studies using a multi-scale analytical framework, with Erica species selected as the model system. Three central findings emerged: first, water availability functions as dominant mechanism mediating climate impacts across all examined contexts; second, while physiological response pathways are generalizable, vulnerability magnitude is context-dependent, reflecting disparities in regional evolutionary histories and endemism levels; and third, climate impacts follow cascading sequence from initial physiological stress through demographic bottlenecks to ecosystem-level changes, potentially creating tipping points where interventions may fail without integrated, multi-scale coordination. This discussion applies the DPSIR framework to organize findings and trace causal pathways from anthropogenic forcing to necessary conservation action, with CFR serving as an illustrative case study whose principles are scalable to Mediterranean-climate biodiversity hotspots globally.

4.1 Synthesis of climate-water-biodiversity pathways: a DPSIR analysis

This DPSIR synthesis integrates evidence from 57 studies to trace causal pathways from global climate drivers to local conservation responses. Only 21% of the reviewed studies reported formal statistical relationships, which precluded the derivation of a meta-analytical threshold. This is a significant methodological consideration that should be acknowledged. Consequently, the quantitative patterns presented throughout this section (e.g., 20%–30% water deficit thresholds) represent convergent evidence synthesized across multiple independent investigations rather than original estimates derived through this review. Geographic bias toward European systems (35% of studies) means these patterns, while operationally useful for conservation planning, require empirical validation for CFR endemic species through targeted manipulative experiments (detailed in Section Research priorities and future directions). With these limitations established, we proceed to apply the DPSIR framework to organize findings and trace pathways from anthropogenic forcing to necessary conservation action, using the Cape Floristic Region as an illustrative case study whose principles are scalable to Mediterranean-climate biodiversity hotspots globally.

Our DPSIR framework operationalizes the assessment of functional conservation areas (Poiani et al., 2000) under climate change conditions. Poiani et al. (2000) defined functional conservation areas through four key ecological attributes: (1) composition and structure of focal ecosystems and species, (2) dominant environmental regimes including natural disturbance, (3) minimum dynamic area needed to maintain ecosystems through disturbance-recovery cycles, and (4) connectivity enabling access to habitats and response to environmental change. The DPSIR framework provides systematic structure for evaluating these attributes under changing conditions: State variables assess composition and structure (population viability, age structure, reproductive output); Pressures evaluate dominant environmental regimes (precipitation patterns, drought frequency, temperature extremes); and Impacts trace consequences for connectivity and minimum dynamic area requirements (demographic bottlenecks limiting dispersal, recruitment failures compromising recolonization capacity). This integration demonstrates how systematic climate-biodiversity synthesis provides empirical foundations for established multi-scale conservation frameworks.

4.1.1 Drivers

Anthropogenic greenhouse gas emissions constitute the fundamental driver, with global temperatures projected to increase 1.5 °C–6.4 °C by 2100 (Intergovernmental Panel on Climate Change (IPCC), 2014), altering precipitation regimes and evapotranspiration demands. These global drivers manifest regionally: Sub-Saharan Africa experienced 0.7 °C warming during the 20th century (Ring et al., 2012; Marcott et al., 2013) with projections of 1.1 °C–6.4 °C by century-end. The Cape Floristic Region shows accelerated warming of 0.1 °C–0.2 °C per decade from 1901–2006 (Haensler et al., 2010), positioning this biodiversity hotspot on a trajectory exceeding the climate conditions under which its endemic flora evolved. Secondary drivers compound direct climate changes. Reduced landscape resilience results from habitat fragmentation and altered hydrological processes caused by land use intensification, urbanization, and water extraction. Beyond just mitigating climate change, these interrelated factors create complex challenges that call for integrated socio-ecological approaches.

4.1.2 Pressures

Climate drivers translate into environmental pressures through multiple pathways. Changes in precipitation patterns, such as decreased total rainfall, increased inter-annual variability, and notable changes in seasonal timing, were found to be the primary climatic pressures at the global Mediterranean scale. Studies from Spain (Prieto et al., 2008; Nogués et al., 2012) and Europe (Llorens et al., 2004) identified summer drought intensification as critical when high evapotranspiration demand coincides with low precipitation. At the Sub-Saharan African scale, pressures manifest through temperature-precipitation interactions. The conditional nature of temperature effects (Jacob et al., 2020), positive under adequate moisture, negative under drought illustrates how pressures emerge from variable combinations rather than single factors. Increased ENSO-associated drought frequency creates episodic but severe pressures exceeding adaptive capacity, even if mean conditions remain tolerable.

Within the Cape Floristic Region, pressures intensify due to the Mediterranean climate, creating predictable summer deficits exacerbated by climate change. Accelerating warming (1980–2000 versus 1950–1970; Warburton et al., 2005), coupled with reduced winter rainfall, creates compound pressures. Critically, convergent evidence from multiple European studies indicates that water deficits of 20%–30% where evapotranspiration demand exceeds water supply represent potential tipping points where synergistic stressors produce non-linear performance declines (Prieto et al., 2008; Llorens et al., 2004). While Prieto et al. (2008) documented this threshold pattern in Spanish E. multiflora, whether CFR endemic species exhibit similar or narrower tolerance ranges remains an empirical question requiring direct investigation.

Within the Cape Floristic Region, pressures intensify due to the Mediterranean climate, creating predictable summer deficits exacerbated by climate change. Accelerating warming (1980–2000 versus 1950–1970; Warburton et al., 2005), coupled with reduced winter rainfall, creates compound pressures. Critically, convergent evidence from multiple European studies indicates that water deficits of 20%–30%, where evapotranspiration demand exceeds water supply, represent potential tipping points where synergistic stressors produce non-linear performance declines (Prieto et al., 2008; Llorens et al., 2004). These thresholds represent initial estimates of boundaries beyond which populations cannot be maintained within their natural ranges of variability (Poiani et al., 2000). Maintaining environmental conditions within this historical variability envelope constitutes a primary conservation objective, as climate-driven exceedance pushes species beyond adaptive capacities evolved under stable conditions. While Prieto et al. (2008) documented this threshold pattern in Spanish E. multiflora, whether CFR endemic species exhibit similar or narrower tolerance ranges remains an empirical question requiring direct investigation.

4.1.3 State

The Cape Floristic Region exemplifies high vulnerability despite exceptional biodiversity. Hosting 9,000+ vascular plant species with 69% endemism (Mittermeier et al., 2011; Manning and Goldblatt, 2012), including approximately 700 endemic Erica species, the region represents a global conservation priority. While the high diversification of Erica species is attributed to rapid speciation over the last 10,000 years under stable climatic conditions, this evolutionary history may have constrained the genetic variation available for rapid adaptation to current environmental change (Table 3). Ecologically, Erica species function as cornerstone taxa (Hitchcock and Rebelo, 2017; Cowling et al., 2015), exerting major influence on soil chemistry, water infiltration rates, and resource availability for associated biota. Current empirical monitoring data substantiate the adverse impacts on plant demographics, particularly illustrating indications of altered phenological patterns in flowering, diminished reproductive output, and a reduction in recruitment frequencies (Dawson et al., 2012; Brown et al., 2016). These indicate climate impacts are present realities requiring immediate response.

East African Afromontane ecosystems present contrasting states. Evolved under greater historical variability, positive growth responses to warming when moisture is adequate (Jacob et al., 2020) suggest some adaptive capacity, though upslope shifts (Kidane et al., 2022) indicate displacement pressures. Topographic complexity provides microrefugia buffering regional trends. European Mediterranean Erica populations exhibit intermediate vulnerability, with broader physiological tolerances from historical climate variability, evidenced by population-specific responses (Llorens et al., 2004; Chamorro et al., 2018).

4.1.4 Impacts

Studies document multiple impact pathways. Elevated thermal conditions augment the process of evapotranspiration, consequently diminishing leaf water potential and instigating the closure of stomata, which results in a reduction of photosynthetic activity (Nogués et al., 2012). Altered precipitation timing affects soil moisture during critical phenological windows, with pre-flowering drought producing 40% flowering reduction in E. multiflora (Prieto et al., 2008). Synthesis of quantitative evidence from European populations indicates that water deficits of 20%–30% trigger non-linear declines, though threshold values vary among populations and among species. These patterns represent fundamental constraints on persistence, but their applicability to CFR endemic taxa requires empirical confirmation.

Demographic impacts cascade from physiological stress. Reduced reproductive output creates recruitment bottlenecks, with germination declining 50%–70% under low moisture (Brown et al., 2016). For fire-adapted seeder species, episodic recruitment failures may generate irreversible population declines. Phenological shifts (2–3-week flowering delays) threaten plant-pollinator synchrony, though direct evidence for CFR pollination mismatches remains limited, a critical research gap. Altered competitive dynamics may favour drought-tolerant species, shifting community composition, with ericaceous cover declines correlated with drying trends (Manzano et al., 2023). Ecosystem-level impacts result from Erica species’ keystone roles (Hitchcock and Rebelo, 2017). Contemporary empirical monitoring data corroborate the adverse impacts on plant populations, particularly illustrating indications of altered phenological flowering periods, diminished reproductive output, and a reduction in recruitment frequencies (Dawson et al., 2012; Brown et al., 2016). These cascades create positive feedback: Erica species loss reduces infiltration → increased runoff → reduced soil moisture, → further Erica species stress. Such feedback suggests potential tipping points where ecosystem shifts become difficult to reverse, possibly when Erica species cover declines below 15%–20% of community composition (Manzano et al., 2023), though this hypothesis requires testing in CFR systems. Impact severity varies with evolutionary and ecological context. European populations tolerate 30%–40% precipitation reductions before comparable impacts (Chamorro et al., 2018), while available evidence from CFR systems suggests threshold responses may occur at 20%–30% reductions (Dawson et al., 2012; Brown et al., 2016), potentially reflecting narrower tolerances from evolution under stable conditions (Table 3). However, direct comparative experiments are needed to confirm these apparent differences in vulnerability.

4.1.5 Responses

Climate-adapted water resource management represents the primary leverage point. The preservation of moisture throughout pivotal phenological intervals, especially the 2–3-month pre-flowering phase, may mitigate the effects of climatic fluctuations. This necessitates the integration of ecological flows into water allocation, protection of riparian corridors and recharge zones, as well as the management of catchments for water retention. In Mediterranean regions with competing demands, explicit allocation for ecosystem functions is necessary but politically challenging. Dynamic conservation planning must anticipate range shifts and facilitate adaptive responses. For montane species, protecting elevational connectivity enables upslope migration (Kidane et al., 2022). However, mountaintop populations face limits, requiring microrefugia identification through climate and topographic modeling. Landscape connectivity is critical for fire-driven fynbos systems, where spatial redistribution depends on habitat permeability increasingly constrained by fragmentation.

Translating conservation strategies into functional landscape design requires explicit consideration of minimum dynamic area, the spatial extent needed to maintain ecosystems through natural disturbance-recovery cycles (Poiani et al., 2000). For CFR Erica communities, this concept is particularly critical given their evolution in fire-driven fynbos systems with 10–30-year fire return intervals. Following Poiani et al. (2000) guideline that functional conservation areas should encompass approximately 50 times the mean disturbance patch size to maintain shifting mosaics of successional stages and given that individual fynbos fires may span hundreds to thousands of hectares, functional landscapes for Erica conservation require 50,000–150,000 ha minimum. This calculation assumes current fire regimes; however, climate change may alter both fire frequencies and intensities, potentially increasing minimum dynamic area requirements. Current protected areas in the CFR should be evaluated against these minimum thresholds, with priority given to: (1) expanding reserves that approach but do not meet minimum dynamic area requirements, (2) establishing functional connectivity among existing reserves to create functional networks (Poiani et al. (2000) capable of sustaining regional-scale processes, and (3) protecting riparian corridors and moisture-retaining landscape positions that serve as recolonization sources following fire events. This spatial planning approach operationalizes Poiani et al. (2000) framework using the water-mediated thresholds and fire ecology evidence synthesized in this review.

Integrated in-situ and ex-situ approaches provide complementary pathways. Seed banking creates insurance against extinctions, particularly for narrow endemics (Pirie et al., 2022). However, maintaining viable in-situ populations preserves evolutionary processes and ecosystem functions. Assisted colonization may become necessary for dispersal-limited species, though ecological risks require evaluation. Population-level climate tolerance variation (Llorens et al., 2004) suggests potential for using adapted genotypes, though translocation guidelines accounting for local adaptation require development. Adaptive management frameworks enable learning as impacts unfold. Given uncertainties in projections and threshold identification, strategies must incorporate monitoring and tracking demographics, phenology, and physiological stress with threshold-based triggers for management adjustment. The conditional and non-linear nature of relationships means static approaches will fail; structured decision-making accounting for uncertainty while enabling adaptation offers greater success potential. Critically, these strategies require multi-scale coordination. Local protected area management must be coupled with regional water policy reform and global climate mitigation participation. No single-scale intervention suffices given nested hierarchies documented here, representing a change in thinking from site-based conservation to integrated landscape and policy-coordinated approaches (Mason et al., 2020; Ranius et al., 2023).

The DPSIR framework offers three advantages addressing gaps identified by Heller and Zavaleta (2009) and Mawdsley et al. (2009). First, unlike narrative reviews organizing thematically (Bellard et al., 2012; Pecl et al., 2017) or functional taxonomies (Mawdsley et al., 2009), DPSIR explicitly structures causal chains from drivers through pressures to state changes to impacts, revealing intervention leverage points. Heller and Zavaleta found 70% of recommendations lacked operational specificity and failed to “distinguish adaptation from climate change impact assessment.” By systematically tracing water-mediated pathways from global forcing (greenhouse gas emissions) through regional alterations (altered precipitation, drought intensification) to local responses (soil moisture deficits; recruitment failure), DPSIR provides the operational specificity they identified as lacking. Second, DPSIR facilitates cross-scale integration that single-level frameworks cannot achieve. While Hannah et al. (2014) microrefugia addresses fine-scale persistence and Peñuelas et al. (2017) synthesis examines regional patterns, neither links processes across scales. DPSIR’s hierarchical structure traces how global drivers generate regional pressures modifying local states, producing species-level impacts necessitating multi-scale responses coordinated from site management through landscape connectivity to policy reform. This addresses the coordination challenges Heller and Zavaleta (2009) identified, operationalizing Xu et al. (2025) demonstration that regional, landscape, and site strategies interact through cascades and policy mechanisms.

Third, DPSIR provides an operational planning structure applicable beyond academic synthesis. By organizing evidence into Driver-Pressure-State-Impact-Response categories, the framework translates into adaptive management cycles: monitoring State variables (soil moisture, flowering, germination) triggers Response adjustments when thresholds approach (e.g., initiating supplemental irrigation when soil moisture falls below 30% during pre-flowering periods), while tracking Pressures (drought intensity, temperature extremes) enables anticipatory management. While Mawdsley et al. identified “improve protected area management” as Strategy 3, they did not specify monitoring variables or intervention thresholds. Our DPSIR demonstrates operationalization: maintain soil moisture >30% during flowering (State threshold), achieve through riparian protection (Pressure modification) and supplementation (Response), evaluate through germination monitoring (Impact assessment). Additionally, DPSIR’s separation of Drivers from Pressures clarifies realistic goals: accepting global temperature increases continue (Driver) while managing water availability (Pressure/State) through riparian protection and ecosystem flows (Responses) to maintain populations within tolerable ranges.

4.2 Generalizable principles versus context-specific vulnerabilities

A central objective was distinguishing ecological principles operating consistently across Mediterranean regions from vulnerabilities arising from CFR’s unique context. Table 4 synthesizes this distinction, revealing that while water-mediated impacts operate through similar physiological pathways globally, vulnerability magnitude is context-dependent. CFR’s exceptional endemism, rapid speciation, and evolution under stable conditions generate heightened sensitivity versus European and East African populations evolved under greater variability.

Table 4

Table 4. Generalizable principles versus context-specific vulnerabilities in Erica species responses to climate-water interactions.

Management strategies based solely on European literature may underestimate CFR vulnerabilities and overestimate adaptive capacity. This extends previous assessments (Bellard et al., 2012; Hannah et al., 2014) by demonstrating that mechanistic pathways differ in quantitative thresholds even when qualitative patterns are consistent: global climate-biodiversity models calibrated on temperate systems may systematically underestimate risks for recently diversified endemic-rich floras, requiring region-specific parameterization. Studies document that climate change affects Erica through direct physiological pathways: elevated temperatures increase evapotranspiration, creating cellular water stress, while altered precipitation modifies soil moisture during critical phases. Nogués et al. (2012) documented 3 °C warming reducing leaf water potential by 15%–20%, triggering stomatal closure and a 25%–30% photosynthesis decline. Prieto et al. (2008) showed pre-flowering drought reduced flower bud formation by 40%, cascading to an estimated 60% seed production decline. Synthesis reveals physiological threshold patterns: growing-season water deficits exceedingly approximately 30% of historical means produce non-linear declines in European populations, though thresholds are population-specific (20% northern, 40% southern: Llorens et al., 2004; Chamorro et al., 2018). Whether CFR endemics exhibit similar patterns or narrower ranges remains an empirical question.

Physiological impacts cascade to demographic consequences through three pathways: reduced reproductive output creates recruitment bottlenecks (50%–70% germination decline: Brown et al., 2016); phenological shifts (2–3-week delays) may desynchronize plant-pollinator interactions; altered competitive dynamics may favour drought-tolerant species (Manzano et al., 2023). Population impacts cascade to ecosystem functioning through Erica keystone roles. Erica loss precipitates deterioration of soil chemical properties and water infiltration capacity, modifies fire regimes, and culminates in microhabitat eradication. These cascades create feedback amplifying impacts, suggesting tipping points beyond which shifts become irreversible, possibly when Erica cover declines below 15%–20% (Hitchcock and Rebelo, 2017).

4.3 Novel contributions as systematic synthesis

This review advances understanding in several ways. First, while previous reviews examined climate impacts broadly (Bellard et al., 2012; Pecl et al., 2017) or focused on specific regions (Peñuelas et al., 2017), none systematically integrated cross-scale evidence to develop mechanistic pathways from global drivers to water-mediated local responses for taxonomically hyper-diverse genera. This mechanistic focus reveals threshold patterns (20%–30% water deficits in European populations), providing operational guidance beyond qualitative risk projections. While these threshold estimates derive primarily from European studies and require validation for endemic CFR species through targeted experiments, they offer practical conservation benchmarks in the absence of species-specific data, a valuable contribution given the urgency of climate adaptation planning.

Second, the explicit DPSIR framework application provides a replicable analytical structure for climate-biodiversity relationships. This review demonstrates how DPSIR links global forcing through regional alterations to local dynamics and management responses, providing a template for similar assessments in other biodiversity hotspots. Third, multi-scale synthesis reveals that generalizable principles must be interpreted within context-specific vulnerability assessments. Table 4’s distinction between universal patterns and CFR-specific sensitivities illustrates how conservation requires both broad physiological understanding and detailed regional evolutionary knowledge. Fourth, quantitative synthesis of relational data (Table 2) addresses a significant deficiency wherein only 21% of original studies provided formalized statistical correlations. While meta-analysis was precluded by methodological heterogeneity, the compiled synthesis provides a foundation for future modeling and hypothesis generation about thresholds requiring empirical testing. Finally, identifying feedback mechanisms and cascading pathways extends prior work on direct effects to encompass indirect pathways through demographic processes and ecosystem interactions. This system’s perspective suggests interventions targeting multiple pathways simultaneously may be necessary to prevent threshold crossings.

4.4 Research priorities and future directions

Several limitations constrain conclusions. European studies represent 35% of publications despite greater CFR diversity, meaning thresholds derived from Spanish E. multiflora may not translate to CFR endemics. Priority research includes coordinated multi-site experiments using standardized protocols enabling formal meta-analysis. The 20%–30% water deficit threshold observed in European populations mandates validation through manipulative experiments spanning distinct species, populations, and life stages in CFR systems. Establishing a stronger empirical basis requires long-term demographic monitoring complemented by microclimate data.

Mechanistically, phenological mismatch concerns a lack of direct CFR empirical confirmation, a critical gap given Erica’s dependence on specialized pollinators. Forthcoming investigations should emphasize quantifying pollinator responses alongside assessing reproductive ramifications of temporal shifts. The genetic basis of climate tolerance remains poorly characterized. Determining whether resilience reflects genetic divergence or phenotypic plasticity is crucial for assessing assisted gene flow viability and forecasting long-term evolutionary trajectories. Integrative genomic methodologies paired with common garden experiments may elucidate adaptive capacity mechanisms. Specific priorities include: (1) experimental manipulations quantifying interactive temperature-precipitation responses across multiple species and populations; (2) long-term demographic monitoring parameterizing population viability models; (3) belowground investigations of mycorrhizal associations and root responses to water stress; (4) genetic analyses identifying climate-adaptive variation aiding assisted migration and restoration while monitoring cumulative threats from fragmentation, invasives, and altered fire regimes.

4.5 Policy recommendations and management priorities

Conservation planning must explicitly recognize water availability as a critical factor affecting species survival (Rolls et al., 2018). Water authorities should develop allocation protocols prioritizing ecological flows during droughts and implement landscape-scale restoration improving water retention (Edwards et al., 2021). Conservation strategies should move beyond static reserve designs to implement dynamic protection, anticipating range shifts and accommodating changing distributions (Ranius et al., 2023), with enhanced landscape connectivity enabling movement across fragmented landscapes (Garden et al., 2015; Littlefield et al., 2019). Conservation strategies such as seed banking, maintaining living collections, and implementing assisted colonization programs function as critical ex-situ insurance mechanisms (Pirie et al., 2022). Successful conservation necessitates seamless integration with in-situ strategies guided by comprehensive species management plans accounting for both present threats and protracted vulnerabilities (Schwartz et al., 2017).

Translating the DPSIR framework and established principles into functional landscape design requires explicit spatial scale considerations. Poiani et al. (2000) minimum dynamic area concept, spatial extent needed to ensure survival or recolonization following disturbance, provides critical operational guidance for CFR Erica conservation. For fire-adapted fynbos with 10–30-year return intervals and fires potentially spanning hundreds of thousands of hectares, Poiani et al. (2000) guideline of conserving areas 50 × mean disturbance patch size suggests functional Erica landscapes require 50,000–150,000 ha minimum to maintain shifting mosaics of fire ages, providing refugia during events and recolonization sources for recovery.

Current CFR protected areas should be evaluated against these minimum dynamic area thresholds, with attention to how climate change may increase spatial requirements through two pathways. First, altered fire regimes, potentially more frequent fires under hotter, drier conditions or fuel load accumulation leading to more intense fires—may increase mean disturbance patch sizes, thereby increasing minimum dynamic area needs proportionally. Second, water stress-induced demographic bottlenecks (reduced germination, limited establishment) may slow post-fire recovery rates, requiring larger spatial buffers ensuring sufficient recolonization sources persist through extended recovery periods. Priority actions: (1) expand reserves approaching but not meeting 50,000-hectare minimum thresholds, particularly those encompassing diverse topographic and moisture gradients; (2) establish functional connectivity among protected areas creating networks (Poiani et al.'s regional-scale conservation unit) enabling species movement across fire-affected landscapes; (3) protect riparian corridors and north-facing slopes as moisture-retaining landscape positions serving as critical recolonization sources; (4) implement adaptive fire management, maintaining natural frequency and intensity ranges while accounting for climate-driven shifts. This spatial planning operationalizes Poiani et al. (2000) functional conservation area typology using water-mediated thresholds, fire ecology evidence, and demographic bottleneck patterns synthesized here, extending Mawdsley et al. (2009) general recommendation to “improve management of existing protected areas” by specifying minimum size requirements based on disturbance ecology.

Furthermore, adaptive conservation requires dedicated funding linked to climate adaptation initiatives (Scherr et al., 2012), and transboundary challenges require coordinated responses transcending political boundaries (Mason et al., 2020), with multi-stakeholder governance forums connecting protected area managers, water utilities, agricultural sectors, and conservation organizations.

4.6 Integration with existing review literature

This multi-scale review demonstrates water availability as the primary mechanism mediating climate impacts on Erica biodiversity, with cascading effects from physiological stress through demographic bottlenecks to ecosystem changes, complementing and extending existing syntheses. While Bellard et al. (2012) and Pecl et al. (2017) documented broad climate-biodiversity relationships, our synthesis reveals water availability as a specific mechanism with quantifiable thresholds (20%–30% deficits in European populations). Bellard et al. (2012) established climate change as a major threat but did not elucidate mechanistic pathways through hydrological changes. Our water-mediated focus provides mechanistic specificity, enabling targeted interventions, knowing that 20%–30% deficits trigger non-linear declines, allows managers to establish monitoring thresholds and implement pre-emptive responses before populations reach tipping points.

Peñuelas et al. (2017) identified drought intensification in Mediterranean forests; we trace cascading pathways revealing intervention leverage points at each organizational level: water supplementation addresses physiological stress, microsite management facilitates demographic transitions, and landscape connectivity maintains ecosystem processes. Previous syntheses identified pressures but did not trace propagation through organizational hierarchies. By demonstrating how water stress cascades from 40% flowering reduction (physiological) through 50%–70% germination decline (demographic) to potentially irreversible ecosystem shifts when Erica cover falls below 15%–20% (functional threshold), our synthesis enables practitioners to intervene at the organizational level most appropriate to their management capacity.

Hannah et al. (2014) microrefugia concept is operationalized through identifying moisture-retaining positions (north-facing slopes, valley bottoms, riparian corridors). Our hydrological synthesis provides practical criteria: areas maintaining soil moisture >30% during summer drought, topographic positions with enhanced infiltration, landscape configurations buffering regional trends enabling practitioners to map microrefugia based on measurable hydrological characteristics rather than abstract “microclimatic heterogeneity. “Building upon foundational frameworks, our synthesis demonstrates how established conservation strategies require context-specific parameterization and cross-scale coordination. Heller and Zavaleta (2009) identified 70% of recommendations lacked operational specificity. We advance by demonstrating water deficit thresholds (20%–30% for European populations, potentially narrower for CFR endemics) provide quantitative targets for implementing general recommendations. Mawdsley et al. (2009) functional taxonomy organized strategies but did not specify implementation thresholds or cross-scale dependencies. Our DPSIR-structured synthesis reveals their Strategy 5 (protect corridors) requires coordination with Strategy 16 (modify water laws) landscape connectivity depends on riparian corridors maintaining adequate moisture requiring regional water governance ensuring ecosystem flows. Xu et al. (2025) demonstration that strategies interact through cascades receives empirical validation: regional water allocation policies enable riparian protection maintaining soil moisture during critical windows, demonstrating nested dependencies they described theoretically.

Critically, our finding that vulnerability is context-dependent despite generalizable mechanisms extends Poiani et al. (2000) foundational work. While Poiani et al. (2000) articulated principles for areas maintaining ecosystems within natural variability, they acknowledged “natural ranges will remain imperfectly known.” By systematically synthesizing water-mediated impacts and documenting threshold patterns (20%–30% deficit), our review provides region-specific, taxon-specific empirical basis for operationalizing their framework. Convergent European evidence establishes an initial estimate of natural variability ranges, while documented narrower tolerances of CFR demonstrate context-dependency of these thresholds. Furthermore, our application of minimum dynamic area to fire-driven fynbos (50,000–150,000 ha) enables evaluating whether current CFR protected areas meet minimum thresholds for maintaining natural disturbance-recovery cycles; an assessment Poiani et al. (2000) called for but could not provide without empirical disturbance data and species-specific vulnerability information.

Furthermore, this synthesis advances understanding of how evolutionary history constrains adaptive capacity. While Yates et al. (2010) called for region-specific parameterization of climate-biodiversity models, our explicit distinction between generalizable principles and context-specific vulnerabilities (Table 4) provides an empirical basis for such parameterization. CFR’s approximately 700 endemic Erica species, products of rapid diversification over 10,000 years under stable conditions, exhibit narrower tolerance ranges than European congeners evolved under greater Quaternary variability. This pattern, consistent physiological mechanisms but divergent vulnerability thresholds has practical implications: conservation strategies successfully applied in European heathlands (where species tolerate 30%–40% deficits) may prove insufficient for CFR without substantial modification accounting for endemic species’ limited adaptive capacity.

For Erica genus specifically, this review extends frameworks established by Fagúndez (2013), Pirie et al. (2022), Pirie et al. (2025). While Fagúndez identified threats to European heathlands, our water-focused synthesis provides mechanistic understanding of how threats propagate: altered fire regimes interact with precipitation changes to modify post-fire soil moisture affecting regeneration success. This mechanistic specificity enables managers to predict interactive effects rather than addressing stressors in isolation. Pirie et al. (2022) outlined ex situ priorities; Pirie et al. (2025) provided systematic gap analysis. This synthesis complements by providing in situ framework: identifying water thresholds for prioritizing at-risk populations, documenting functional diversity as insurance mechanism, establishing minimum dynamic area requirements for fire-adapted systems. Integration of ex situ insurance with our in-situ water-mediated management framework provides comprehensive guidance spanning germplasm preservation through wild population management to landscape-scale ecosystem conservation. Additionally, Pirie et al. (2024) demonstration of spatial decoupling between species richness and phylogenetic diversity confirms planning must incorporate evolutionary dimensions; our framework emphasizes functional diversity recognizing rapid diversification under stable conditions constrains adaptive capacity, requiring protective rather than adaptive conservation strategies for many CFR endemics.

While this review focuses primarily on ecological mechanisms, Feliciano et al. (2025) analysis of climate impacts on small-scale versus large-scale farmers underscores that conservation strategies must be developed in conjunction with livelihood considerations. Where human communities depend on ecosystem services provided by biodiversity-rich landscapes like CFR, including water regulation, pollination, and genetic resources, conservation planning requires integrating social dimensions. The water-mediated framework provides ecological foundations, but successful implementation depends on governance structures equitably distributing adaptation costs and benefits across stakeholder groups. This remains a critical research priority for translating ecological understanding into socially sustainable conservation outcomes. This synthesis advances methodological approaches through three innovations: explicit DPSIR structure enabling systematic organization (response to Heller and Zavaleta’s call); multi-scale framework creating a two-dimensional matrix revealing geographic and biological hierarchies; systematic distinction between generalizable and context-specific patterns (Table 4), providing a template for other hotspots. However, limitations constrain conclusions: only 21% of studies reported statistical relationships precluding meta-analysis; geographic bias toward Europe (35%) limits generalization to CFR endemics; qualitative synthesis restricts effect size precision.

Critical knowledge gaps persist. First, 20%–30% water deficit threshold requires validation through coordinated CFR experiments. Second, phenological mismatch lacks direct CFR confirmation critical gap given Erica’s dependence on specialized pollinators. Third, genetic basis of tolerance remains poorly characterized. Fourth, interactive effects among water stress, altered fire, invasives, and fragmentation require integrated assessment. Finally, social-ecological dimensions (Feliciano et al., 2025) demand integration ensuring strategies are socially equitable and politically feasible. Looking forward, the multi-scale, mechanistically grounded framework provides both conceptual foundation and practical guidance for evidence-based planning. By explicitly linking established frameworks (Poiani et al., 2000; Heller and Zavaleta, 2009; Mawdsley et al., 2009; Xu et al., 2025) with empirical water-mediated thresholds and context-specific vulnerability assessments, this synthesis demonstrates how systematic evidence integration operationalizes theoretical principles. The challenge ahead lies in implementing coordinated multi-scale responses with sufficient speed and scope to maintain CFR Erica communities within natural ranges of variability before irreversible tipping points are crossed.

5 Conclusion

This multi-scale systematic review demonstrates water availability as the primary mechanism mediating climate change impacts on Erica biodiversity across Mediterranean-climate regions, with cascading effects from physiological stress through demographic bottlenecks to ecosystem functional changes. Through the synthesis of 57 studies organized within the DPSIR framework, three central contributions emerge, advancing both theoretical understanding and practical conservation application.

First, while physiological response mechanisms operate consistently across Mediterranean regions water deficits consistently reduce flowering, germination, and establishment vulnerability magnitude is fundamentally context-dependent. CFR’s approximately 700 endemic Erica species, products of rapid recent diversification over 10,000 years under stable climatic conditions, exhibit narrower tolerance ranges than European congeners evolved under greater Quaternary climate variability. This finding challenges the application of global climate-biodiversity models calibrated on temperate systems to endemic-rich floras, demonstrating the necessity for region-specific parameterization in megadiverse regions.

Second, quantitative synthesis reveals threshold patterns where water deficits of 20%–30% relative to historical conditions distinguish resilient from vulnerable populations. These thresholds provide operational benchmarks for conservation planning and represent empirical quantification of natural ranges of variability (Poiani et al., 2000) for precipitation regimes supporting Erica populations. Documentation of cascading impacts 40% flowering reduction cascading to 50%–70% germination decline, potentially culminating in irreversible ecosystem state changes when Erica cover falls below 15%–20% underscores the urgency of implementing comprehensive conservation responses before irreversible ecosystem state changes occur. The non-linear, threshold-dependent nature of these relationships means incremental climate changes may produce disproportionate biodiversity impacts once critical thresholds are exceeded.

Third, the DPSIR framework application demonstrates how systematic evidence synthesis informs multi-scale conservation coordination. By explicitly tracing causal pathways from global atmospheric forcing through regional hydrological alterations to local biodiversity responses, this review provides a replicable template for climate-biodiversity assessments in other hotspots. The framework’s hierarchical structure reveals intervention leverage points at multiple scales: regional water policy reform modifies pressures before they trigger state changes; landscape-scale riparian corridor protection maintains states within tolerable ranges; site-scale water supplementation provides a direct response to state deficits; demographic monitoring tracks impacts, enabling adaptive response adjustment. This multi-level intervention strategy operationalizes theoretical principles while providing spatial planning guidance through minimum dynamic area requirements (50,000–150,000 ha for fire-driven fynbos systems).

The systematic distinction between generalizable principles and context-specific vulnerabilities offers practical guidance for adapting conservation strategies. Universal patterns include water availability positively correlates with reproductive success across all Mediterranean Erica; temperature effects are conditional on water availability (positive when wet, negative when dry); summer drought impacts exceed other seasonal periods due to evapotranspiration peaks. Context-specific vulnerabilities include CFR species showing narrower precipitation tolerance ranges than European congeners; limited thermal plasticity from evolution under a stable Mediterranean regime; disproportionate keystone functional roles (15%–20% of CFR community composition versus 2%–5% in European heathlands), meaning Erica loss triggers cascading ecosystem impacts.

Safeguarding CFR’s unique evolutionary heritage requires integrated action coordinated across spatial scales and governance levels. Climate-adapted water resource management ensuring ecosystem flows during critical phenological windows particularly the 2–3-month pre-flowering period when soil moisture deficits most severely impact reproductive output represents the primary conservation leverage point. Landscape-scale conservation planning must incorporate microrefugia identification through hydrological modeling (protecting north-facing slopes, valley bottoms, riparian corridors maintaining >30% soil moisture saturation during summer drought) and elevational connectivity enabling upslope range shifts. Protected area design should meet minimum dynamic area thresholds (50,000–150,000 ha based on fire ecology) with functional connectivity among reserves creating regional-scale networks supporting recolonization following disturbance. Adaptive management frameworks with threshold-based monitoring triggers enable responsive strategy adjustment as impacts unfold.

The convergence of exceptional endemism (700 endemic Erica species representing a globally unique evolutionary experiment), narrow physiological tolerances, limited adaptive capacity, and accelerating climate change (0.1 °C–0.2 °C per decade warming, projected summer drought intensification) positions CFR among the world’s most vulnerable biodiversity hotspots. This vulnerability is compounded by the socio-economic value of Erica species their documented ethnomedicinal applications and phytochemical diversity mean biodiversity loss eliminates not only evolutionary heritage and ecosystem functions but also valuable biological resources. Feedback mechanisms whereby Erica loss reduces soil infiltration, increasing runoff and reducing moisture availability, create positive feedback, accelerating further loss that may become infeasible to reverse.

In conclusion, this systematic review demonstrates that effective conservation of Mediterranean-climate biodiversity hotspots under accelerating climate change requires: (1) mechanistic understanding of water as the primary mediating pathway enabling targeted interventions; (2) recognition that vulnerability magnitude is context-dependent despite universal physiological mechanisms; (3) cross-scale coordination linking global atmospheric forcing through regional water governance to local population management; (4) threshold-based adaptive management responding pre-emptively to approaching tipping points; and (5) integration of in situ population management with ex situ germplasm conservation. The DPSIR-structured, multi-scale framework developed here provides both conceptual foundation and operational guidance for implementing these principles, offering a replicable template for climate-biodiversity assessments and conservation planning in other biodiversity hotspots facing water-mediated climate impacts.

Data availability statement

The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding authors.

Author contributions

ZM: Conceptualization, Data curation, Formal Analysis, Investigation, Methodology, Project administration, Resources, Software, Validation, Writing – original draft, Writing – review and editing. MS: Conceptualization, Formal Analysis, Investigation, Methodology, Resources, Software, Supervision, Validation, Writing – review and editing.

Funding

The author(s) declared that financial support was not received for this work and/or its publication.

Acknowledgements

I gratefully acknowledge the instrumental supervision and advice provided by Mulala Danny Simatele, whose constructive feedback was vital to this paper. Institutional support was provided by the University of the Witwatersrand library, which furnished necessary access to scholarly databases and resources. The successful completion of this project was also critically dependent on my sustained commitment and tenacity throughout the challenging academic process. The extensive investment of time during the research and writing stages was instrumental, facilitating both the conclusion of this work and substantial personal and intellectual development.

Conflict of interest

The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Generative AI statement

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