Fungal Phenology as an Event-Scale Indicator of Climate Variability and Ecosystem Function

Srishti Verma^1*, Samay Tirkey², Shobhana Ramteke³, Manas Kanti Deb⁴

^1,3,4 School of Studies in Environmental Science, Pt. Ravishankar Shukla
University, Raipur, Chhattisgarh, India

²School of Studies in Life Sciences, Pt. Ravishankar Shukla

University, Raipur, Chhattisgarh, India

Abstract

 Fungi play a critical role in ecosystem processes, but the dynamic responses of fungi to climate change are still poorly understood. This review explores fungal phenology as a sensitive, event-scale bioindicator of environmental change. Unlike plants, the fruiting of fungi is directly controlled by external conditions, particularly by the soil moisture and temperature. Therefore, fruiting events are frequently triggered shortly after rain and are representative of short-term microclimatic conditions. Now evidence is emerging that fungal phenology is already changing in response to climate change. The fruit season is lengthening. Timing is shifting. Patterns are also becoming more unpredictable. These shifts are mediated by temperature-precipitation interactions and are heterogeneous in space and function. Changes in fruiting also affect species composition and community structure. Fungal phenology is an underutilized ecological indicator that can provide near real-time signals of soil status. It complements plant-based indicators as it captures fine scale variability often missed. Changes in fungal activity also have broader ecological implications. They can alter trophic interactions, nutrient cycling and carbon dynamics. There is more interest, but there are still big gaps. Most studies have focused on temperate regions and aboveground observations are little integrated with belowground processes. The review emphasizes the need for standardization, wider geographical coverage and integration of molecular and ecological data.

Keywords: Bioindicators; Climate change; Ecosystem functioning; Fungal phenology; Soil microclimate

1.     Introduction

Fungi play a central role in terrestrial ecosystems, yet their responses to climate variability remain relatively understudied. As decomposers and symbiotic partners, they regulate nutrient cycling, organic matter turnover, and plant productivity through saprotrophic and mycorrhizal interactions (Boddy, 2016; Baldrian et al., 2022). In doing so, they contribute directly to ecosystem stability by mediating carbon flux, and the nutrient exchange between soil and vegetation (Crowther et al., 2016; Rillig et al., 2002). Despite of this, the fungal dynamics, especially their temporal responses, have received far less attention than plant and animal phenology.

Phenology is widely used to detect ecological responses to climate change. Shifts such as earlier flowering, altered migration, and longer growing seasons are now well documented. Fungi, however, follow a different logic. Fruiting body development depends directly on environmental conditions, particularly soil moisture and temperature, rather than internally buffered physiological controls (Kauserud et al., 2008; Gange et al., 2011; Büntgen et al., 2012; Boddy et al., 2014). As a result, fungal phenology often reflects the current environmental conditions rather than seasonal averages. For instance, saprotrophic species such as Pleurotus ostreatus and Coprinus comatus can fruit rapidly after rainfall, whereas ectomycorrhizal fungi like Boletus edulis or Cantharellus cibarius show responses shaped by both climate and host plant dynamics. This close environmental coupling gives fungal fruiting particular ecological relevance. Fruiting bodies are short-lived structures that appear only when specific microclimatic thresholds are met, within few days of precipitation or temperature shifts (Egli et al., 2010; Karavani et al., 2018). In this sense, fungi operate as event-scale indicators, meaning biological indicators that respond rapidly to short-term environmental events such as rainfall pulses, transient moisture availability, or abrupt temperature shifts, thereby capturing fine temporal variability in soil conditions that broader seasonal indicators may overlook. Moisture pulses act as primary triggers, while the temperature defines the window of response.

Evidence now shows that fungal phenology is already shifting under climate change. Long-term datasets reveal changes in timing, duration, and variability of fruiting, including the delayed autumn peaks, earlier onset in some systems, and extended fruiting seasons (Kauserud et al., 2008; Kauserud et al., 2010; Andrew et al., 2018). These patterns reflect interacting effects of temperature and precipitation, though the responses differ across regions and functional groups (Büntgen et al., 2012; Vitasse et al., 2021). Changes in fruiting intensity and species composition further suggest the ongoing reorganization of fungal communities (Andrew et al., 2016; Morera et al., 2022). These characteristics support the view of fungi as high-resolution indicators of environmental variability. Unlike the conventional bioindicators that integrate long-term conditions, fungal phenology provides near real-time signals of soil microclimate dynamics (Diez et al., 2013; Halme et al., 2012). Yet this potential remains underused, partly due to the limited geographic coverage and methodological inconsistency.

Fungi are also embedded within ecological networks. Many organisms depend on fungal fruiting bodies as seasonal resources, and shifts in timing can disrupt these relationships, leading to trophic mismatches (Crowther et al., 2016; Tedersoo et al., 2022). Changes in fungal activity can also influence the nutrient cycling, carbon dynamics, and soil structure, amplifying the ecological consequences of climate change (Baldrian et al., 2022).

Despite increasing attention, major gaps remain. Most studies focus on temperate regions, with limited data from tropical and monsoon-driven systems where fungal diversity and climate sensitivity are high (Andrew et al., 2016; Tedersoo et al., 2022). In addition, the methodological inconsistencies and limited integration of belowground processes restrict predictive understanding.

Although fungal phenology has increasingly been linked with climate change, most existing studies primarily describe seasonal shifts in fruiting patterns rather than evaluating fungi as event-scale ecological indicators. Current reviews also tend to focus either on fungal productivity or broad climate responses, with limited emphasis on the mechanistic coupling between short-term microclimatic variability, soil processes, and ecosystem functioning. In addition, integration between aboveground phenological observations and belowground ecological dynamics remains insufficient, particularly across tropical and monsoon-driven systems. This review addresses these gaps by synthesizing fungal phenology within the framework of event-scale environmental sensitivity and ecosystem-level climate responses.

In this context, here we review our current knowledge of fungal phenology as a climate-responsive process. More specifically it considers (a) the biological basis of fungal sensitivity to environmental drivers (b) empirical evidence for climate-driven phenological changes (c) the potential for fungi as high-resolution indicators of climate variability and (d) the ecological consequences of altered fruiting dynamics. By synthesizing these perspectives, we contend that fungal phenology is a powerful but underappreciated framework for understanding ecosystem responses to climate change (Fig. 1).  

 2.     Biological Basis of Fungal Sensitivity to Climate Variability

Fungal phenology reflects direct exposure to environmental conditions rather than internally regulated timing. Fruiting depends on external cues, primarily the soil moisture, temperature, and substrate availability without the buffering seen in plants (Boddy et al., 2014; Büntgen et al., 2012). This leads to responses that are rapid, threshold-driven, and often short-lived.

Figure 1. Climate variability influences the fungal phenology via soil microclimate. Threshold-driven responses to moisture and temperature may alter fruiting timing, duration, and frequency, with cascading effects on trophic interactions and nutrient cycling.

Soil moisture plays a central role. Fruiting typically follows rainfall events that rehydrate the mycelial network and trigger primordia formation (Egli et al., 2010; Karavani et al., 2018). Species such as Coprinus comatus and Agaricus bisporus often respond quickly to such moisture pulses. Continuity of moisture is critical; brief or interrupted wet periods can lead to weak or irregular fruiting, which contributes to variability under changing precipitation regimes. Temperature shapes the conditions under which these responses occur. Metabolic activity and developmental rates vary across the species-specific thermal ranges (Boddy, 2016). Warmer conditions can shift fruiting patterns, including the delayed autumn peaks or earlier spring emergence, however only where the moisture remains sufficient (Kauserud et al., 2008; Kauserud et al., 2010; Büntgen et al., 2012). Moisture and temperature therefore act together rather than independently. Substrate and carbon availability add further complexity. Saprotrophic fungi depend on organic matter inputs and decomposition dynamics (Boddy, 2016; Morera et al., 2022), whereas mycorrhizal fungi rely on host-derived carbon. In species such as Boletus edulis and Amanita muscaria, environmental stress which affects the host plants can limit carbon allocation and reduce fruiting (Rillig et al., 2002).

A defining feature of fungi is their structural simplicity. Hyphae remain directly exposed to surrounding conditions, and allowing rapid responses to environmental change (Boddy et al., 2014). Fruiting may occur quickly under favorable conditions and cease just as rapidly when conditions deteriorate. Responses differ across the functional groups. Saprotrophic fungi generally track the moisture pulses closely, while mycorrhizal fungi exhibit more complex patterns which is shaped by both climate and host physiology (Andrew et al., 2016; Tedersoo et al., 2022). This variability limits species-level generalization and supports the use of functional group approaches. Climate variability amplifies these dynamics. The Drought, intense rainfall, and temperature extremes can all disrupt fungal activity (Karavani et al., 2018). Fruiting patterns may shift from predictable seasonal events to irregular or multi-peak occurrences. Overall, the fungal phenology operates near environmental thresholds, with the responses triggered only when the conditions align. This sensitivity underlies both the vulnerability to climate change and the potential as a fine-scale ecological indicator.

 3.     Evidence of Climate-Driven Shifts in Fungal Phenology

Evidence from long-term datasets shows clear shifts in fungal fruiting patterns under the changing climate conditions. Changes are not limited to timing; they extend to duration, frequency, and the intensity of fruiting. Across the regions, one consistent signal emerges that, ‘the fungal phenology is becoming more variable and less predictable’.

Temporal shifts are widely reported. In temperate systems, autumn fruiting often occurs later, while some spring-fruiting species appear earlier, effectively stretching the fruiting season (Kauserud et al., 2008; Kauserud et al., 2010; Andrew et al., 2018). Rather than a simple shift forward or backward, the pattern reflects an expansion of the reproductive window. This extended activity aligns with the warmer conditions that prolong periods suitable for fungal growth. Temperature and precipitation jointly shape these changes. Temperature influences the metabolic activity and seasonal windows, while precipitation, especially soil moisture controls the initiation of fruiting (Büntgen et al., 2012; Karavani et al., 2018). In many ecosystems, rainfall events act as immediate triggers, which links the fruiting more closely to short-term weather patterns than to the long-term averages (Egli et al., 2010). Shifts in rainfall distribution, including drought periods and irregular precipitation, therefore lead to disrupted or uneven fruiting cycles.

Regional differences remain pronounced. The temperate regions, particularly Europe, show extended fruiting seasons and delayed autumn peaks (Kauserud et al., 2008; Andrew et al., 2018). Mediterranean systems display stronger dependence on rainfall timing, with marked fluctuations in productivity across years (Morera et al., 2022; Karavani et al., 2018). In mountain ecosystems, elevation gradients interact with warming trends, and alters both the distribution and phenological timing (Vitasse et al., 2021). Data from tropical and monsoon-driven regions remain limited, though these systems likely exhibit strong rainfall-driven responses. Beyond the timing, the changes in fruiting intensity and community composition are increasingly reported. Climate variability can suppress moisture-sensitive species while favoring more tolerant taxa, and leading to a shift in dominance and functional structure (Andrew et al., 2016; Baldrian et al., 2022). These changes indicate not only phenological adjustment but also community reorganization. A notable emerging pattern lies in increased interannual variability. The fruiting events may occur in multiple pulses within a season or fail entirely in certain years. This irregularity reflects growing influence of extreme events such as droughts, heatwaves, and intense rainfall, that disrupt stable environmental conditions (Karavani et al., 2018). As a result, fungal phenology shifts from predictable seasonal cycles to more fragmented and event-driven dynamics.

Advances in data availability have strengthened these conclusions. Long-term monitoring plots provide detailed temporal trends, while herbarium records and citizen science datasets extend observations across the larger spatial scales (Diez et al., 2013; Halme et al., 2012). When it is combined with climate data, these sources reveal consistent links between fungal activity and environmental variability, despite the differences in methodology. Overall, the current evidence points to a transition from stable, seasonally structured fruiting towards the more variable and climate-sensitive patterns. Changes in timing, duration, and community composition all reflect the close coupling between fungi and environmental conditions, which reinforce their value as indicators of climate variability.

4.     Fungi as indicators

Fungal phenology offers a different way to read environmental change-closer to real-time conditions than to seasonal averages. Fruiting occurs only when specific combinations of moisture and temperature align. Which turns fungal activity into a direct expression of soil microclimate (Büntgen et al., 2012; Boddy et al., 2014). In contrast to many biological indicators that reflect accumulated conditions, the fungi register short-term environmental states.

This sensitivity originates from a threshold-dependent behavior. Fruiting does not follow a gradual response curve; it appears once environmental limits are crossed. The soil moisture acts as the immediate trigger, while the temperature defines the window within which this response becomes possible. The result is a system that responds to events such as rainfall pulses, brief favorable periods, rather than some long-term trends. Observed fruiting, or its absence, therefore provides a direct signal of underlying soil conditions (Egli, 2011; Karavani et al., 2018). A key advantage lies in the integration of belowground processes. The soil microclimate remains spatially variable and difficult to measure directly. Mycelial networks, already embedded within this environment, respond to these variations and translate them into the visible fruiting bodies. In this way, the fungi act as biological proxies for soil conditions that are otherwise hard to capture with conventional measurements (Diez et al., 2013; Halme et al., 2012).

Comparison with plant phenology highlights this distinction. Plant responses often reflect seasonal progression and are moderated by the internal regulation. Fungal responses occur faster and track short-term fluctuations more closely. This difference does not replace plant-based indicators but complements them, and adding resolution at a finer temporal scale. Where the plants indicate seasonal change, the fungi indicate immediate environmental states. The indicator value of fungi also depends on a scale. Individual observations reflect local conditions, while the aggregated datasets reveal broader patterns. Large-scale records derived from the monitoring plots, herbarium collections, and citizen science that allow links between the fungal phenology and climate variability to emerge across regions (Diez et al., 2013; Halme et al., 2012). These datasets support the development of predictive relationships, however the consistency in data collection remains a limitation.

Several constraints shape interpretation. Responses vary among species and functional groups, detection depends on the observation frequency, and most datasets remain concentrated in the temperate regions (Andrew et al., 2016; Tedersoo et al., 2022). In addition, fruiting represents only the visible phase of fungal activity, while much of the system operates below the ground surface. Despite of these limitations, fungal phenology provides a useful framework for the environmental monitoring. Its strength lies not in replacing existing indicators, but in adding a layer that captures short-term variability and the soil-level processes. With improved standardization and broader geographic coverage, the fungi could serve as practical indicators of climate variability, ecosystem condition, and the environmental stress.

5.     Ecological Consequences of Shifting Fungal Phenology

Changes in fungal phenology extend beyond timing. They alter ecological interactions and core ecosystem processes. Since the fungi occupy the central positions in food webs and biogeochemical cycles, even the small shifts in fruiting patterns can propagate through multiple trophic and functional pathways (Boddy et al., 2014; Crowther et al., 2016).

One of the most immediate effects appears in trophic interactions. Fungal fruiting bodies serve as seasonal resources for a wide range of organisms, including insects, rodents, and other mycophagous fauna (Tedersoo et al., 2022; Boddy, 2016). Many of these organisms rely on the predictable fruiting periods. Shifts in timing or duration can disrupt this synchrony, and leading to the mismatches between the resource availability and the consumer demand. Such mismatches may reduce survival or reproductive success in dependent species, and particularly where life cycles are tightly linked to the fungal availability (Crowther et al., 2016; Tedersoo et al., 2022). These disruptions are especially pronounced in insect–fungus interactions. Many insects develop directly within fruiting bodies, and making their life cycles closely aligned with fungal phenology. Altered timing or reduced fruiting can limit suitable substrates, and affecting the larval development and the community composition. Since these insects also contribute to spore dispersal and the decomposition processes, changes at this level may feed back into the fungal dynamics. Beyond the trophic effects, shifts in phenology influence the nutrient cycling and carbon dynamics. Fungi regulate decomposition and nutrient release, which link aboveground inputs to soil processes. Changes in fungal activity can modify the decomposition rates, alter the nutrient availability, and influence the soil carbon storage (Baldrian et al., 2022; Crowther et al., 2016). Periods of reduced activity, such as drought, may slow the organic matter breakdown, while favorable conditions may accelerate it. These fluctuations introduce variability into processes that are typically more stable.

At the community level, climate-driven phenological shifts often coincide with the changes in species composition. The moisture-sensitive species may decline under drier conditions, while tolerant or generalist taxa increase in dominance (Andrew et al., 2016; Morera et al., 2022). Such shifts alter functional balance within the fungal communities, with consequences for decomposition pathways, symbiotic interactions, and the ecosystem stability. A broader pattern emerges in the form of increasing variability. The fruiting events may occur in multiple pulses, shift unpredictably, or fail entirely in certain years. This irregularity reduces the reliability of seasonal cues for organisms that depend on fungi and can destabilize the ecological networks. Systems that rely on synchronized interactions become particularly vulnerable under these conditions.

These changes also affect ecosystem services. Fungi contribute to the soil fertility, carbon regulation, and food resources, including the wild edible mushrooms. Variability in the fruiting patterns can influence both the ecological functioning and the human uses, particularly in regions where mushrooms support livelihoods and traditional practices (Procházka et al., 2023; Guo and Xu, 2025). Overall, shifting the fungal phenology reshapes ecosystems through a combination of trophic disruption, altered nutrient cycling, and community reorganization. The effects are not isolated; they interact and reinforce one another, and reflecting the central role of fungi in linking climate variability to ecosystem function.

6.     Future Directions and Research Priorities

Progress in fungal phenology research has been substantial, hwoever the field remains uneven in scope, data quality, and integration. The strongest limitation is geographic bias. Most evidence comes from temperate regions, while tropical and monsoon-driven systems, where fungal diversity and climate sensitivity are high, remain poorly represented (Andrew et al., 2016; Tedersoo et al., 2022). This restricts global generalization and limits understanding of rainfall-driven dynamics that likely to dominate large parts of the world.

A second constraint lies in the data structure and comparability. Studies differ in how the phenology is defined and measured first appearance, peak fruiting, duration, or presence/absence that makes synthesis difficult (Diez et al., 2013). In addition, the fungal fruiting is inherently ephemeral and spatially patchy, which leads to strong detection bias. Long-term monitoring provides high-quality data but remains geographically limited; herbarium records extend historical coverage but reflect collector bias; citizen science expands spatial scale but introduces variability in identification and sampling effort (Halme et al., 2012). These datasets are valuable, though none alone provides a complete picture.

A deeper gap concerns the gap in connection between the aboveground observations and belowground processes. The fruiting bodies represent only a transient phase, while most fungal activity occurs as mycelium in soil. Molecular approaches, including eDNA and metabarcoding, have begun to reveal the hidden diversity and activity patterns (Andrew et al., 2016; Baldrian et al., 2022). Integration between these approaches and phenological observations remains limited, and constraining interpretation of how visible fruiting relates to underlying ecosystem processes. Ecological understanding also remains incomplete. While trophic mismatch is recognized conceptually, the empirical links between fungal phenology and consumer dynamics are scarce. Responses likely to vary across systems and taxa, however coordinated studies tracking both fungi and dependent organisms remain rare (Crowther et al., 2016; Tedersoo et al., 2022). Similarly, most analyses focus on mean climatic trends, while the climate extremes and event-scale dynamics, droughts, intense rainfall, heatwaves receive less attention despite of their strong influence on fungal responses (Karavani et al., 2018).

These limitations point toward clear priorities. First, to expand the monitoring into underrepresented regions, particularly the monsoon-driven ecosystems, will be essential for capturing global patterns. Second, to standardize the phenological metrics onset, peak, duration, and intensity would enable a meaningful comparison across the studies. Third, to integrate the data types, field observations, historical records, molecular datasets, and high-resolution environmental measurements offers a path toward more robust inference. Advances in statistical modeling, such as hierarchical and process-based approaches, provide the tools for linking these datasets and improving predictive capacity (Diez et al., 2013).

Technological developments can accelerate this progress. Automated detection systems, image-based monitoring, and high-throughput sequencing expand both the temporal resolution and spatial coverage, while reducing the observer bias (Barauskas et al., 2022). At the same time, a shift toward the functional trait-based approaches can reduce the complexity by focusing on ecological strategies rather than the individual species (Andrew et al., 2016). Looking forward, the field needs to move beyond the description toward prediction and application. The Incorporation of fungal phenology into ecosystem and climate models will improve representation of decomposition, carbon cycling, and soil processes (Crowther et al., 2016; Baldrian et al., 2022). At a practical level, fungi offer potential as indicators of the soil moisture dynamics, ecosystem stress, and the resource availability, particularly in the regions where livelihoods depend on wild mushrooms (Procházka et al., 2023; Guo and Xu, 2025). Overall, the advancement in fungal phenology requires a shift from fragmented observations to integrated frameworks. Linking the climate drivers, soil processes, fungal responses, and ecological interactions will position fungi not only as sensitive indicators, but also as active components of climate ecosystem feedbacks.

7.     Conclusion

Fungal phenology offers a distinct and underused perspective on the climate and ecosystem interactions. Unlike the plant-based indicators that reflect seasonal trends, the fungal fruiting tracks immediate environmental conditions, particularly at the soil level. This close coupling with moisture and temperature makes fungi sensitive to both gradual change and short-term variability, which position them as practical indicators of shifting environmental states.

Current evidence shows that fungal phenology is already changing fruiting periods are extending, timing is shifting, and patterns are becoming less predictable. These changes are not isolated. They influence the trophic interactions, nutrient cycling, and broader ecosystem functioning, often in the ways that remain insufficiently understood. At the same time, responses vary across the regions and functional groups, which highlight the limits of generalization and the need for more context-specific understanding. A key challenge moving forward lies in connecting what is observed above the ground with the processes occurring below the ground. The fruiting bodies provide visible signals, but they represent only a part of the fungal activity. Integrating the phenological observations with molecular and environmental data will be necessary to fully interpret these dynamics. Expanding the research beyond temperate systems, particularly into monsoon-driven and tropical regions, is equally important.

Fungi are unlikely to replace existing ecological indicators, but they can complement them in the meaningful ways. Their strength lies in capturing short-term variability and soil-level processes that are otherwise difficult to detect. With improved data integration, the standardized approaches, and broader geographic coverage, the fungal phenology could contribute not only to the climate change research but also to practical monitoring of ecosystem condition. Rather than treating fungi as a secondary component of ecological studies, there is a value in placing them more centrally within the climate and ecosystem frameworks. Doing so may reveal the patterns and processes that remain overlooked, particularly in a system where the environmental change is rapid and uneven.

 Acknowledgments

All the authors are thankful to Pt. Ravishankar Shukla University, Raipur, Chhattisgarh, India, for providing infrastructure and laboratory facilities.

 Conflict of interest

Dr. Srishti Verma, Dr. Samay Tirkey, Dr. Shobhana Ramteke and Prof. Manas Kanti Deb declare that they have no conflict of interest.

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