ABSTRACT
Reservoirs are increasingly recognized as important sources of greenhouse gases (GHGs), yet the temporal scales and environmental drivers governing CO2 and CH4 flux variability remain poorly constrained. This is particularly the case for Mediterranean systems, which are subject to strong hydrological variability. Here, we present and analyze year‐round, high‐frequency eddy covariance measurements of CO2 and CH4 fluxes in a Mediterranean reservoir over two years with contrasting hydrological conditions, explicitly evaluating variability at diurnal, multiday, and seasonal timescales. The reservoir acted as a net source of both gases in both years, but CH4 fluxes exhibited substantially greater temporal variability than CO2 fluxes. The seasonal component dominated CH4 flux variability (> 75%), with annual areal fluxes and reservoir‐wide emissions 44% higher during the drier year due to an earlier onset and longer duration of the high‐emission summer period. Near‐sediment temperature and water‐column depth were the primary predictors of seasonal CH4 emissions, reflecting temperature controls on methanogenesis and hydrological regulation of ebullition. In contrast, CO2 flux variability was dominated by diel processes, with wind speed emerging as the main predictor of short‐term emissions, while water depth and surface temperature regulated seasonal variability. For both gases, chlorophyll‐a was an important predictor at multiday timescales. Our results demonstrate that the temporal variability of GHG fluxes from Mediterranean reservoirs is highly sensitive to hydrologically mediated changes in lake water levels, particularly for methane. Under climate change, GHG emissions could increase significantly in Mediterranean regions, and potentially in other seasonally dry regions worldwide, as a result of projected trends towards stronger eutrophication and increases in drought frequency and duration. Our study underscores the value of long‐term, high‐frequency observations to improve emission estimates and to better represent reservoir processes in regional and global carbon cycle assessments.
Keywords: Eddy covariance, eutrophic reservoir, greenhouse gases, hydrological variability, Mediterranean climate, wind‐forced gas exchange
We monitored greenhouse gas emissions from a Mediterranean reservoir over 2 years that differed markedly in water availability. Methane emissions were much higher during the drier year, largely because lower water levels and warmer bottom waters favored methane production and release, while carbon dioxide emissions were less affected. Our results highlight the sensitivity of reservoir methane emissions to drought and their potential to increase under future climate change.

1. Introduction
Inland freshwater ecosystems, despite occupying a small fraction of the Earth's surface (Messager et al. 2016), play an outsized role in the global carbon cycle (Tranvik et al. 2009), contributing substantially to global atmospheric CO2 and CH4 emissions (Rosentreter et al. 2021; DelSontro et al. 2018; Raymond et al. 2013). In lakes and reservoirs, although CO2 emission rates are roughly four times higher than CH4 emission rates, CH4 is the dominant greenhouse gas (GHG) in terms of climate forcing due to its much higher global warming potential, accounting for approximately 75% of total CO2‐equivalent emissions in these lentic systems (DelSontro et al. 2018; Deemer et al. 2016). Recent global bottom‐up assessments indicate that more than 40% of global methane emissions originate from aquatic ecosystems, with lakes and reservoirs alone accounting for over 10% (Rosentreter et al. 2021). Large uncertainty is still inherent in these estimates, as observational data coverage is insufficient—especially with respect to episodic fluxes, the complete annual cycle, and spatial coverage (e.g., Rodríguez‐Velasco et al. 2024; Johnson et al. 2022, 2021; Deemer and Holgerson 2021; Deemer et al. 2016).
Air–water CO2 and CH4 fluxes integrate the effects of a wide range of physical, biological, and, particularly in the case of CO2, chemical processes operating across multiple temporal scales within a year. CO2 fluxes reflect the balance between metabolic processes such as net ecosystem production and allochthonous carbon inputs (Lapierre et al. 2013; McDonald et al. 2013; Cole et al. 2007), carbonate equilibrium and calcite precipitation (Many et al. 2024), catchment lithology (León‐Palmero, Morales‐Baquero, and Reche 2020), as well as physical forcing associated with stratification, mixing, and synoptic weather variability (e.g., Reed et al. 2018; Liu et al. 2016). These processes can interact with trophic dynamics, including phytoplankton–zooplankton cycles and the development of algal blooms, further modulating CO2 exchange at diel to seasonal scales (Ouyang et al. 2017). In contrast, methane emissions are primarily governed by sedimentary methanogenesis under anoxic conditions and by oxidation (Bastviken et al. 2004; Mayr et al. 2020) and/or further production in the water column (Ordóñez et al. 2023; León‐Palmero, Contreras‐Ruiz, et al. 2020; Tang et al. 2014), with fluxes strongly shaped by physical transport and release mechanisms. Seasonal stratification and turnover, ice formation, hydrological variability, and episodic pressure‐ and temperature‐driven ebullition events exert strong control on CH4 emissions (Ustinov et al. 2025; Jammet et al. 2015; Yvon‐Durocher et al. 2014; Bastviken et al. 2004). Nutrient loading and eutrophication further modulate methane production and release by enhancing labile organic matter supply for methanogenesis (Martínez‐García et al. 2024; Grasset et al. 2018; West et al. 2015).
Beyond intra‐annual variability, another source of uncertainty in estimates of lake and reservoir CO2 and CH4 emissions arises from interannual variability in both flux magnitudes and the relative importance of their controlling processes. Year‐to‐year changes in climate forcing, hydrology, and ecosystem state can alter the balance between physical transport, biogeochemical production and consumption, and biological activity, potentially leading to shifts in the dominant drivers of gas exchange. This interannual variability is particularly relevant in the Mediterranean region (roughly 27°–47° N, 15° W–40° E) and more generally, in regions with a Mediterranean climate, which are characterized by a pronounced warm and dry season and a highly uneven distribution of precipitation during the remainder of the year. As a result, lakes and reservoirs in this region commonly experience large seasonal fluctuations in water temperature and water level, with strong implications for stratification, mixing dynamics, and sediment–water interactions. In addition, the Mediterranean climate exhibits marked interannual and multi‐decadal variability in precipitation (e.g., Esteban‐Parra et al. 2022), an increasing trend in evaporative water losses due to rising air temperatures (e.g., Esteban‐Parra et al. 2022), and widespread anthropogenic alterations of catchments. Reservoirs are the dominant lentic ecosystems in this region (e.g., Lehner and Döll 2004), and water‐level drawdown during summer is often intensified to meet irrigation and domestic water demands, further amplifying hydrological variability.
Despite the recognized importance of lakes and reservoirs in global carbon budgets and the substantial intra‐ and interannual variability of CO2 and CH4 fluxes, much of our understanding and assessment is still based on discrete measurements of fluxes conducted with flux chambers or gas traps that are deployed intermittently, or indirect gas‐transfer models (e.g., Deemer et al. 2016; Wik et al. 2013; Bastviken et al. 2004). While these approaches have provided valuable insights, they are often limited in temporal resolution, spatial coverage, and their ability to capture episodic or scale‐dependent flux dynamics. In contrast, eddy covariance (EC) systems enable continuous, long‐term, and spatially integrated non‐invasive measurements of air–water gas exchange (e.g., Baldocchi 2014), offering a unique opportunity to quantify carbon fluxes across multiple temporal scales. To date, only a small subset of EC studies have examined the temporal patterns or environmental drivers of CO2 and/or CH4 fluxes in lakes or reservoirs over complete annual cycles or across multiple years, whether for CO2 (Golub et al. 2023; Scholz et al. 2021; Reed et al. 2018; Ouyang et al. 2017; Liu et al. 2016; Mammarella et al. 2015; Shao et al. 2015; Huotari et al. 2011), CH4 (Waldo et al. 2021; Iwata et al. 2020; Taoka et al. 2020) or both gases (Hounshell et al. 2023; Spank et al. 2023; Eugster et al. 2020; Jammet et al. 2017). Only four of these studies were conducted in temperate reservoirs (Hounshell et al. 2023; Spank et al. 2023; Waldo et al. 2021; Liu et al. 2016), and only two of these (Hounshell et al. 2023; Spank et al. 2023) measured CO2 and CH4 concurrently. Moreover, none of these studies explicitly examined reservoir carbon fluxes under hydrologically contrasting years, which are common in Mediterranean climates. Consequently, our understanding of how ecosystem transitions and contrasting or altered hydrological regimes regulate reservoir CO2 and CH4 fluxes—and their underlying drivers—remains limited. Here, we use EC measurements from a shallow Mediterranean reservoir spanning two hydrologically contrasting years—a dry year with low precipitation and reduced water storage, and a wet year—to identify the dominant temporal scales of CO2 and CH4 flux variability, determine the key environmental predictors at each scale and their interannual shifts in importance, and quantify the influence of hydrological variability on reservoir carbon flux dynamics.
2. Methods
2.1. Study Site
Cubillas Reservoir is a small, shallow, polymictic eutrophic reservoir located in southeastern Spain (37.276° N, −3.671° E). The reservoir is primarily used for irrigation and recreation. It has been in operation since 1956 and drains a 647 km2 calcareous, predominantly agricultural watershed (e.g., León‐Palmero, Morales‐Baquero, and Reche 2020), with Cubillas River as the main inflow. The reservoir has a surface area of 1.94 km2, and its initial total capacity was 19 Hm3. Current capacity is generally much lower and averaged (reference period 1991–2020) 13 ± 4 Hm3, with water levels at 638 ± 3 m.a.s.l, resulting in annual maximum depths near the dam of 13.6 ± 2.8 m (Figure 1a). Within a year, the reservoir experiences an average water‐level drawdown of 5.4 ± 2.1 m (Figure 1a), with maximum levels typically occurring in April–May and minimum levels at the beginning of each hydrological year (October). For this study, we use data collected in 2022 and 2024, which represent contrasting hydrological conditions. Although both years were warm (positive air temperature anomalies; Figure 1b), Year 2022 (hereafter the dry year) was characterized by a cumulative precipitation of 447 mm (close to the historical average, Figure 1c), a maximum water depth of 8 m, and an average inflow rate of 0.34 m3 s−1. In contrast, 2024 (hereafter the wet year) had a cumulative precipitation of 532 mm (positive anomaly, Figure 1c), a maximum reservoir depth of 9.4 m, and inflow rates of 0.42 m3 s−1. Although 2023 exhibited a larger negative precipitation anomaly than 2022, EC malfunction throughout the entire summer prevented temporal analyses of carbon fluxes for that year.
FIGURE 1.

Historical meteorological and hydrological anomalies for the Cubillas Reservoir from 1980 to 2024. (a) Annual maximum reservoir water depth and annual water‐level drawdown, (b) mean annual air temperature (T air), and (c) cumulative annual precipitation and number of wet days (i.e., days with > 1 mm of precipitation). Anomalies were calculated relative to the 1991–2020 reference period. Historical air temperature and precipitation data were derived by interpolating all available observations from the State Meteorological Agency (AEMET), the Automatic Hydrological Information System (SAIH), and the Andalusian Agroclimatic Information Network (RIA) meteorological stations, as in Herrero (2007). Maximum reservoir depth was derived from reservoir bathymetry, historical water‐level records, and historical changes in hypsographic curves provided by the Guadalquivir Hydrographic Confederation. Grey shaded areas indicate the 2 years studied.
2.2. Data Collection
We deployed a 5 × 5 m aluminum floating platform at 37.278° N, −3.672° E (Figure S1) in December 2021. The platform is anchored to the reservoir sediment using four 300 kg concrete blocks, deployed at its four vertices and positioned 40–50 m from the platform. At ≈10 m from the platform, each of the four mooring ropes has an additional 30‐kg concrete block, which acts as an additional anchorage during low‐water level conditions and limits the platform's motion and rotation during water level changes. The platform holds an accelerometer (WTGAHRS2, Wit‐Motion) to track platform rotation angles (yaw, pitch, roll). The accelerometer measured at a rate of 1 Hz until March 2023, when it was increased to 10 Hz.
The observation platform was equipped with an EC measurement system installed at a height of 3.34 m. The EC system includes a 3D ultrasonic anemometer (Windmaster, Gill) to measure wind speed and direction, as well as two open‐path infrared gas analyzers to measure CH4 (LI‐7700, LI‐COR), and CO2 and water vapor (LI‐7500DS, LI‐COR) in the ambient air. The EC system records data at a rate of 10 Hz. Since installation, the system also provides data on air temperature and relative humidity every minute, and starting on May 5, 2022, it also records incoming and reflected shortwave radiation and incoming and outgoing longwave radiation (CNR4 net radiometer, Kipp & Zonen) with the same frequency. All the flux and meteorological data collected by the EC system were integrated into the SmartFlux system (LI‐COR).
Additional 5‐min meteorological data, including atmospheric pressure, wind speed and direction (2D WindSonic M, Gill), and incoming photosynthetically active radiation PAR (LI‐190R, LI‐COR), were also available at the platform. In addition, one surface mooring and one bottom mooring, both fixed to the platform, provided measurements of other environmental variables. For this study, we used: (i) 1‐min water temperature records from three high‐frequency thermistors built by a local manufacturer (Edrónica), calibrated to a precision of 0.01°C, and located 0.5 and 1 m below the lake surface and 0.5 m above the lake sediment; and (ii) 1‐min records of dissolved oxygen DO concentration collected with optical sensors (TrioOS) at 1 m depth and 0.5 m above the lake sediment. To convert height above the sediment to depth below the surface, we also installed a piezoresistive level probe (36XW, Keller) at a height of 0.5 m above the lake sediment. Hanging at 0.5 m below the lake surface, a fluorometer (nanoFlu, TrioOS) provided data on chlorophyll‐a concentration every 5 min. To avoid photoquenching effects (e.g., Rousso et al. 2021), we only used nighttime values. We further retained only chlorophyll‐a measurements collected after sensor cleaning; before June 2024, cleaning was performed manually on a regular weekly basis, while from June 2024, an automatic wiper enabled daily quality‐controlled values. Hourly data on water level, reservoir volume and inflow and outflow discharges are available from the Guadalquivir Hydrographic Confederation website (https://www.chguadalquivir.es/saih/Inicio.aspx; last access on 29 January 2026).
2.3. EC CH4 and CO2 Flux Calculations
We obtained 30‐min CH4 and CO2 fluxes from the raw 10 Hz EC data using the EddyPro v.7.0.9 software (LI‐COR). Flux processing followed established international standards and protocols, including spike detection and removal (Vickers and Mahrt 1997), double rotation for sensor tilt correction (Wilczak et al. 2001), covariance maximization with default settings for time‐lag compensation between the anemometer and the gas analyzers, and block‐averaging to extract turbulent fluctuations from the time series. Corrections applied included the Webb–Pearman–Leuning (WPL) density correction for open‐path analyzers (Webb et al. 1980), as well as spectral corrections for high‐frequency (Moncrieff et al. 1997) and low‐frequency (Moncrieff et al. 2004) signal attenuation. For all the fluxes, we calculated quality check flags (Mauder and Foken 2004) to test for developed turbulent conditions and stationarity. We selected only 30‐min fluxes with the best quality flags (flag values = 0). We further filtered fluxes based on the automatic gain control (AGC) values of the open‐path gas analyzers to ensure optical cleanliness. Specifically, we retained periods with AGC ≥ 70% for the CO2 gas analyzer and AGC ≥ 20% for the CH4 gas analyzer.
We conducted a second analysis to evaluate platform rotation on EC flux estimate errors using the raw 10 Hz EC data for 2024 for which yaw, pitch and roll rotational angles were also available at 10 Hz. Prior to the flux calculations with EddyPro, we corrected the raw 3D ultrasonic anemometer data for sensor tilt using the measured yaw, pitch and roll angles (e.g., Spank et al. 2023), and we compared the results with the unrotated 30‐min fluxes. Pitch and roll rotation of the platform led to non‐significant differences between rotated and non‐rotated signals (Figure S2), demonstrating that rotational effects along the horizontal x and y axes could be discarded. The wind direction estimated by the EC systems was corrected for yaw angle platform rotation before footprint calculations. We calculated the sampling area of the EC tower with the 2D flux footprint model of Kljun et al. (2015). We retained all 30‐min CO2 and CH4 flux measurements for which the wind was coming from directions where 80% of the footprint (80% contour line) was over the water at the lowest water level reached in a given year. Flux measurements were also removed when wind direction indicated potential influence from the dock or from a transient island that emerges in the northern sector of the reservoir during periods of low water level. This translated into removing wind directions between −36° and 50° in 2022 and −25° and 50° in 2024 (Figure S1). After quality control and footprint analysis, we retained 23% of the 30‐min CO2 and 36% of CH4 fluxes in 2022, and 25% of CO2 fluxes and 33% of CH4 fluxes in 2024. These percentages fall within the range reported for EC measurements over lakes and reservoirs (e.g., Hounshell et al. 2023; Erkkilä et al. 2018). We filled gaps in the 30‐min flux time series for CO2 and CH4 using the REddyProc online tool (Wutzler et al. 2018; https://bgc.iwww.mpg.de/5622399/REddyProc). Gaps remaining in the series (less than 1.5%) were filled through linear interpolation. Time series of gas fluxes were averaged into hourly and daily fluxes.
2.4. Wavelet Analysis
We partitioned the original hourly time series of gap‐filled CH4 fluxes into diffusive and ebullitive components using the wavelet analysis approach proposed by Iwata et al. (2018), based on local scalar similarity and dissimilarity between CH4 concentration and air temperature, respectively. The hourly time series of gap‐filled CO2 and CH4 fluxes (including the partitioned ebullitive and diffusive components) measured by the EC system were decomposed into contributions at different temporal scales using the maximal overlap discrete wavelet transform (MODWT) (e.g., Golub et al. 2023; Knox et al. 2021; Sturtevant et al. 2016). The MODWT was applied using a Symlet wavelet of order 8 (sym8) (Sturtevant et al. 2016). The wavelet coefficients were computed using the MATLAB function modwt, and the corresponding multiresolution analysis (MRA) components were reconstructed using modwtmra. This procedure decomposed the original signal into a set of scale‐dependent detail components and a residual smooth component, whose sum equals the original time series.
Given the hourly sampling interval, each MODWT level j corresponds to a specific range of temporal scales of 2j h. We aggregated the reconstructed components into three broader time‐scale bands (e.g., Knox et al. 2021): (i) a diurnal component, corresponding to periods between approximately 8 and 32 h; (ii) a multiday component (synoptic to sub‐seasonal variability), corresponding to periods between approximately 3 and 43 days; (iii) a seasonal component, corresponding to periods longer than approximately 43 days. We performed aggregation by summing the reconstructed wavelet components whose characteristic periods fell within each band (Figures S3–S6). We calculated the variance of each aggregated component and expressed it as a fraction of the total variance of the original interpolated time series. In this manner, we could quantify the relative contribution of diurnal, multiday, and seasonal variability to the overall variance of CO2 and CH4 fluxes. Due to the redundancy and non‐orthogonality of the MODWT, as well as boundary effects, the sum of variances across multiresolution components does not necessarily equal the total variance; therefore, variance fractions should be interpreted as approximate contributions.
2.5. Attribution Study and Statistical Analysis
To attribute variability at each temporal scale to environmental and biogeochemical predictors, we fitted Random Forest (hereafter RF) regression models independently to the original fluxes and to each decomposed flux component. We used a common set of 15 potential predictor variables (Figure S7; see Section 2.2 for measurement resolution), including air temperature (T air), near‐surface (T surf) and bottom water temperature (T bot), thermal stratification strength (ΔT = T surf−T bot), atmospheric pressure (P atm), water column depth (Depth), the temporal gradient of total pressure (atmospheric plus hydrostatic) (ΔP tot/Δt), wind speed, bottom‐water oxygen concentration (O2,bot), surface chlorophyll‐a concentration (Chl‐a), inflow discharge as a surrogate for allochthonous nutrients and carbon inputs, incoming shortwave radiation (SWR), net surface buoyancy flux (B 0,net) as a precursor for convective turbulence, net ecosystem production (NEP) and gross primary production (GPP). NEP and GPP were derived from oxygen concentrations following the diel dissolved oxygen method (Staehr et al. 2010), based on a mass‐balance approach that partitions diel O2 dynamics into biological production, respiration, and air–water gas exchange. Although Random Forests are relatively robust to predictor collinearity for prediction, strongly correlated variables can bias permutation‐based importance metrics; therefore, we reduced collinearity before fitting the model. We assessed collinearity among predictors using Spearman's rank correlations. Because air temperature and surface and bottom water temperatures were strongly correlated (|R| > 0.8, Figure S8), only bottom temperature was retained for CH4 flux models and surface temperature for CO2 flux models based on their expected physical relevance. SWR was also strongly correlated with B 0,net; therefore, B 0,net was retained given the known influence of convection on diffusive gas fluxes (e.g., Eugster et al. 2003; MacIntyre et al. 2010). Precipitation over the reservoir was not included as a predictor variable. Although rainfall events have been linked to short‐term variability in CH4 ebullitive emissions in some systems (e.g., Niu et al. 2025), rainfall in Mediterranean regions typically occurs outside the summer high‐ebullition season.
We implemented RF models with identical hyperparameters across gases, years, and temporal scales, including a fixed number of trees (300) and permutation‐based out‐of‐bag (OOB) predictor importance. We fitted the models separately for each year to assess interannual variability in driver importance. We evaluated predictor importance using the increase in OOB prediction error following random permutation of each predictor, and we interpreted the results in terms of relative rankings and grouped process contributions rather than absolute importance values. To assess the robustness of predictor importance estimates to the stochastic nature of the algorithm (bootstrap resampling and random predictor selection), each analysis was repeated 50 times using different random seeds. The reported importance values correspond to the mean across runs. NEP and GPP were originally derived at daily resolution from dissolved oxygen measurements and linearly interpolated to the hourly time step to match the temporal resolution of the EC flux data. Chlorophyll‐a concentrations were available at weekly resolution until June 2024 and at daily resolution thereafter and were similarly interpolated to an hourly time step. We applied these interpolations solely to align predictors with the flux time series and did not introduce additional high‐frequency variability; consequently, the influence of these predictors is expected to be expressed primarily at multiday to seasonal time scales. The remaining predictors were collected at sub‐hourly resolution (see Section 2.2) and aggregated to hourly resolution by averaging.
To test for differences between daytime and nighttime conditions, we analyzed the original hourly methane (CH4) and carbon dioxide (CO2) fluxes. We defined daytime and nighttime using photosynthetically active radiation (PAR): hours with PAR ≥ 10 μmol photons m−2 s−1 were classified as daytime, and hours with PAR < 10 μmol photons m−2 s−1 as nighttime. To account for the temporal autocorrelation of hourly measurements, fluxes were aggregated to daily mean values for daytime and nighttime periods. This produced a dataset with one daytime and one nighttime flux estimate per day. We tested differences between daytime and nighttime fluxes using a linear mixed‐effects model, with daytime/nighttime as a fixed effect and day as a random intercept to account for repeated measurements within days. We fitted models using restricted maximum likelihood (REML). Statistical significance of diel differences was assessed using the fixed‐effect coefficient for day‐night, with α = 0.05. Conclusions were not sensitive to moderate changes in the PAR threshold used to define daytime conditions.
3. Results
3.1. Methane and Carbon Dioxide Emissions at Different Temporal Scales
The reservoir acted as a net source of both CO2 and CH4 independently of the year considered (wet/dry). Daily carbon fluxes varied markedly between years, with the greatest interannual differences observed in CH4 fluxes (Figure 2). During the dry year, CH4 fluxes averaged 16.83 ± 20.48 mmol m−2 d−1 and exhibited significant seasonal changes, with minimum fluxes on the order of 10−1 mmol m−2 d−1 in winter and early spring, and maximum fluxes reaching up to 73 mmol m−2 d−1 in summer. In the wet year, mean CH4 fluxes decreased to 11.44 ± 20.11 mmol m−2 d−1 while maintaining a similar seasonal range of variability. This reduction in average fluxes during the wet year is primarily attributable to a delayed onset of the summer high‐flux period, which began on June 28—almost 2 months later than in the dry year, when it started on May 1 (Figure 2a). As a result, the cumulative annual flux in the dry year (5.9 mol m−2 year−1) was 44% higher than that in the wet year (4.1 mol m−2 year−1) (Figure 2b). Assuming that the EC footprint is representative of the entire reservoir, cumulative CH4 emissions amounted to 8.05 × 106 and 5.6 × 106 mol year−1 during the dry and wet years, respectively (Figure S9). Thus, reservoir‐wide emissions were also 44% higher during the dry year, despite seasonal differences in reservoir surface area of up to 20 ha. Ebullition dominated over diffusion, accounting for 70% and 75% of the cumulative areal fluxes in the dry and wet years, respectively (Figure 2b).
FIGURE 2.

Time and cumulative series of CH4 and CO2 fluxes. EC diurnal averaged (a, c) and 30‐min cumulative (b, d) gap‐filled CH4 (a, b) and CO2 (c, d) fluxes for the two studied years. Cumulative fluxes in (b) also include the partition of the total flux into ebullition and diffusion. Shaded areas in (a, c) indicate ±1 standard deviation. X‐axis ticks mark the first day of each month.
CO2 fluxes exhibited weaker seasonality than CH4 fluxes. During the dry year, CO2 fluxes averaged 40.95 ± 57.18 mmol m−2 d−1, with maximum values of 166 mmol m−2 d−1 occurring in late spring (Figure 2c). In the wet year, the seasonal maximum occurred earlier, in early spring. However, no significant interannual variability was observed, as mean fluxes were similar (41.06 ± 54.15 mmol m−2 d−1, or 14.9 mol m−2 year−1; see Figure 2d). Reservoir‐wide cumulative CO2 emissions amounted to 20.54 × 106 and 22.65 × 106 mol year−1 during the dry and wet years, respectively (Figure S9), indicating that total CO2 emissions were approximately 10% higher during the wet year due to the larger reservoir surface area.
Seasonality dominates the temporal variability of CH4 fluxes, with the decomposed seasonal component explaining more than 75% of the variance in the original signal in both years (Figure 3). Diurnal variability is the second most important contributor, accounting for 14% and 9% of the total variance during the dry and wet years, respectively (Figure 3). Variability at the multiday scale explains less than 6% of the total variance in both years. In contrast, diurnal variability dominates the temporal variability of CO2 fluxes (Figure 3), with the decomposed diurnal component explaining 42% and 48% of the total variance in the original signal during the dry and wet years, respectively. The contribution of the multiday scale differed between years, explaining 27% of the variance in the dry year but only 17% in the wet year. Seasonality accounted for a smaller share, explaining 11%–13% of the total variance.
FIGURE 3.

Contribution of the different temporal scales to the total variance of the hourly gas flux signals. Results derived from the discrete wavelet transform and multiresolution analysis of the hourly gap‐filled CH4 and CO2 flux time series.
3.2. Predictor Variables for CH4 and CO2 Fluxes
In both years, bottom water temperature (T bot), the temporal gradient in total pressure (ΔP tot/Δt), wind speed, and water column depth were the four major predictors of the original CH4 signal (R 2 > 0.95; Figure 4a; Figures S10–S11), in the RF analysis. During the wet year, wind speed surpassed bottom water temperature in importance, probably due to the shorter duration of the summer high‐flux (ebullitive) period compared to that of the dry year (Figure 2a). When the signal was decomposed into its temporal components, T bot clearly emerged as the dominant driver of seasonality, followed by water column depth (R 2 ≥ 0.999; Figure 4b; Figures S10–S11). Multiday variability was well explained by variations in T bot and water column depth (R 2 ≥ 0.970; Figure 4c; Figures S10–S11). Finally, diurnal variability was best predicted by wind speed, T bot, ΔP tot/Δt, and additional ancillary variables, with some differences between years (R 2 > 0.88; Figure 4d; Figures S10–S11).
FIGURE 4.

Random forest predictor importance for CH4 (a–d) and CO2 (e–h) fluxes during the dry and wet years. Random Forest regression models were applied separately to the original flux time series (a, e) and to the decomposed flux components obtained via maximal overlap discrete wavelet transform (MODWT) and multiresolution analysis: seasonal (b, f), multiday (c, g), and diurnal (d, h) signals. Each Random Forest analysis was repeated 50 times using different random seeds to assess model stochasticity. Bars represent mean predictor importance across runs, and black horizontal lines indicate ±1 standard deviation.
For CO2 fluxes, wind speed was the main RF predictor of both the original CO2 signal and the decomposed diurnal components in both years (Figure 4e,h), explaining between 40% and 53% of the variability in these signals (Table S1). Seasonal variability was primarily explained by water column depth and surface temperature T surf (R 2 > 0.999; Figure 4f; Figures S12–S13). Water column depth alone explained 75% and 97% of the variability in the seasonal CO2 flux signals during the dry and wet years, respectively (Table S1). Multiday variability was explained by temporal variations in water column depth, T surf, Chl‐a concentrations, and GPP (R 2 > 0.970). Water column depth, together with Chl‐a, was more influential during the dry year, whereas Chl‐a, GPP, and T surf better predicted multiday variability during the wet year (Figure 4g).
Bottom water temperature alone explained more than 80% and more than 90% of the variability in the original and seasonal CH4 flux signals, respectively (Table S1). The sensitivity of methane emissions to temperature can be described using an exponential Arrhenius‐type function (e.g., Yvon‐Durocher et al. 2014):
| (1) |
where is the methane flux at temperature T in K, is the methane flux at 20°C, E A (eV) is the activation energy, and k b is the Boltzmann constant (8.62 × 10−5 eV K−1). The activation energy was estimated by fitting Equation (1) to the daily‐averaged CH4 flux measurements from both years combined (Figure 5a), yielding E A = 1.89 ± 0.01 eV (R 2 = 0.9989). The sensitivity to water temperature was approximately 70% higher during the wet year (E A = 2.60 ± 0.02 eV) than during the dry year (E A = 1.53 ± 0.02 eV). When separating ebullition and diffusion, the fitted activation energy for the ebullitive fluxes was E A = 2.02 ± 0.02 eV (R 2 = 0.9979), twice the activation energy for the diffusive fluxes (E A = 1.00 ± 0.01 eV, R 2 = 0.9980; Figure S14). The seasonal pattern exhibited hysteresis in the relationship between water temperature and methane emissions, with higher emissions at a given temperature during the second half of the year (Figure 5a). This hysteretic behavior is explained by the second most important driver, the temporal evolution of water column depth (Figure 4b): as water depth—and thus hydrostatic pressure—decreases over the course of the summer season, the release of methane as bubbles is facilitated (Figure 5a). Chlorophyll‐a (Chl‐a) was the third most important seasonal predictor in the RF analysis (Figure 4b).
FIGURE 5.

Hysteretic relationship between water temperature and methane fluxes, and diurnal decomposed ebullitive and diffusive signals. (a) Bottom water temperature vs. methane flux. Red and blue dots represent the original daily‐averaged methane fluxes plotted against the measured daily‐averaged water temperature at 0.5 m above the lake sediment. The filled scattered circles indicate the relationship between the decomposed seasonal bottom‐water temperature T bot and methane flux signals, with colors denoting the corresponding seasonally decomposed water‐column depth at the EC platform location. For visualization purposes, the seasonal relationships for the wet year are shown in a separate inset. Black arrows indicate the counterclockwise direction of the hysteresis over the annual cycle. The black dashed line shows the best‐fit Arrhenius relationship based on the two‐year dataset, with the corresponding activation energy (E A ) indicated. The gray shaded area around the Arrhenius fit represents the 95% confidence interval. (b) Diurnal decomposed ebullitive signal and the ΔP tot/Δt signal, and (c) diurnal decomposed diffusive signal and wind speed for a selected summer period during the dry year. Vertical red lines in (b,c) indicate the time of flux peaks. Time is in UTC.
The magnitude of the diurnal component of the ebullitive fluxes varied with ΔP tot /Δt (Figure 5b; Figure S15b), with higher ebullition rates occurring during rapid decreases in total pressure in the afternoon. The diurnal cycle commonly exhibited a bimodal pattern, with a secondary peak in the early morning, corresponding to a secondary drop in total pressure. In contrast, the decomposed diurnal pattern of diffusive methane fluxes followed the diurnal cycle of wind speed over the reservoir, with peak values occurring in the afternoon (Figure 5c; Figure S15a). The linear mixed‐effects model applied to year‐round original hourly methane fluxes, using daytime/nighttime as a fixed effect, indicated that methane fluxes were significantly higher during the daytime. On average, daytime fluxes were 50% and 20% higher than nighttime fluxes during the dry and wet years, respectively (Table 1; Figure S16).
TABLE 1.
Linear mixed‐effects model results for day‐night differences in CH4 and CO2 fluxes.
| Year | Gas | Effect | Estimate a (μmol m−2 s−1) | 95% CI (μmol m−2 s−1) | p | % change a |
|---|---|---|---|---|---|---|
| 2022 | CH4 | Day vs. Night | 0.0759 | [0.065, 0.0867] | 2.84e‐38 c | 49 |
| 2022 | CO2 | Day vs. Night | 0.0610 | [0.0243, 0.0977] | 0.001 c | 14 |
| 2024 | CH4 | Day vs. Night | 0.0240 | [0.0165, 0.0314] | 5.30e‐10 c | 20 |
| 2024 | CO2 | Day vs. Night | −0.0388 | [−0.077, −0.0006] | 0.046 b | −8 |
The estimate indicates the difference between daytime and nighttime fluxes. The percentage change measures the differences between the mean daytime and nighttime fluxes divided by the mean nighttime fluxes.
Significant at 95% level.
Significant at 99% level.
Unlike CH4 fluxes, CO2 fluxes showed a weak dependence on temperature, with low activation energies of ≈0.03 eV (Figure 6a,b). During the dry year, the decomposed seasonal CO2 flux signal decoupled from the seasonal surface water temperature signal (Figure 6c), and a clear hysteretic pattern emerged in the relationship between water temperature and CO2 fluxes (Figure 6b), with lower emissions at a given temperature during the second half of the year as the reservoir water level decreased. As with diffusive methane fluxes, the decomposed diurnal pattern of CO2 fluxes showed a clear synchronous relationship with wind speed (Figure 6d). The linear mixed‐effects model applied to year‐round hourly CO2 fluxes, with daytime/nighttime included as a fixed effect, indicated contrasting patterns between years. During the dry year, CO2 fluxes were significantly higher during the daytime, by an average of 14%. In contrast, during the wet year, CO2 fluxes were significantly lower during the daytime, by an average of 8% (Table 1; Figure S16).
FIGURE 6.

Sensitivity of CO2 fluxes to temperature and diurnal emissions. (a,b) Surface water temperature versus CO2 flux during the (a) dry and (b) wet years. Red and blue dots represent the original daily‐averaged CO2 fluxes plotted against the measured diurnal‐averaged water temperature at 0.5 m below the lake surface. Filled scatter symbols show the relationship between the seasonally decomposed surface water temperature (T surf) and CO2 flux signals, with colors indicating the corresponding seasonally decomposed water‐column depth at the EC platform location. Black arrows in (b) indicate the clockwise direction of hysteresis over the annual cycle. The black dashed lines in (a, b) show the best‐fit Arrhenius relationships for each year, with the corresponding activation energies (E A ) indicated. The gray shaded areas around the Arrhenius fits represent the 95% confidence intervals. (c) Seasonally decomposed T surf and CO2 flux signals. (d) Diurnal decomposed CO2 flux signal and wind speed for a selected summer period during the wet year. Vertical blue lines indicate the timing of flux peaks. Time is in UTC.
4. Discussion
Average methane emissions from Cubillas fall within the range reported for tropical reservoirs (e.g., Prairie et al. 2021) and lie within the upper decile of emission rates reported for reservoirs globally (e.g., Prairie et al. 2021; Deemer et al. 2016). Strong seasonal patterns in methane fluxes as found in Cubillas have been observed in shallow temperate, boreal, and subarctic lakes and reservoirs (e.g., Waldo et al. 2021; Jansen et al. 2020; Podgrajsek et al. 2016), as well as in wetlands (e.g., Knox et al. 2019; Sturtevant et al. 2016). Seasonal dependencies between methane fluxes and air, water, or sediment temperature have also been documented in reservoirs (e.g., Kim et al. 2025), lakes (e.g., Jansen et al. 2020; Jammet et al. 2017; Wik et al. 2014), and wetlands (e.g., Knox et al. 2021) worldwide. These patterns ultimately reflect the temperature dependence of methanogens' metabolic activity during the decomposition of organic matter in anoxic sediments (e.g., Yvon‐Durocher et al. 2014). The relationship between CH4 emissions and temperature is commonly exponential and described using an Arrhenius‐type equation (e.g., Iwata et al. 2020; Jansen et al. 2020; Natchimuthu et al. 2016). The exponential temperature dependence observed at Cubillas is consistent with previous year‐round measurements in the reservoir using a flux chamber coupled to a gas analyzer (Rodríguez‐Velasco et al. 2024), and the estimated activation energies for ebullition (E A ≈2 eV) and diffusion (E A ≈1 eV) fall within the range of values reported in earlier studies (Praetzel et al. 2021; Jansen et al. 2020; Aben et al. 2017; DelSontro et al. 2016).
Here, we further show that CH4 emissions exhibit a pronounced hysteretic pattern, with higher emissions at a given temperature during the second half of the year (Figure 5a). Such counterclockwise hysteretic behavior is commonly observed in wetlands (Chang et al. 2021), but has been rarely reported in lakes or reservoirs (Jansen et al. 2020). This hysteretic pattern has been attributed to seasonal differences in microbial abundance and/or substrate availability and characteristics in the sediment (e.g., Chang et al. 2020; Updegraff et al. 1998), both of which influence methane production. In Cubillas, chlorophyll‐a (Chl‐a) arose as the third most important predictor of the seasonal component of CH4 fluxes in the dry year. Chl‐a concentrations increase as the summer season progresses, potentially enhancing CH4 production as this readily degradable material is deposited in the sediments (e.g., Martínez‐García et al. 2024; Grasset et al. 2018; West et al. 2015, 2012), thereby reinforcing the hysteretic effect. However, water column depth—and thus hydrostatic pressure—arises in our RF analysis as the major driver controlling CH4 flux hysteresis in Cubillas. Together, near‐sediment temperature and water column depth explain more than 99% of the seasonal variability in CH4 fluxes. Reduced hydrostatic pressure lowers the threshold for bubble formation and release, increasing the contribution of ebullition to the overall fluxes, which also bypasses water‐column oxidation. Consequently, methane fluxes are higher at a given temperature later in the season under shallower water depths, producing the observed hysteresis in the methane flux‐temperature relationship. By showing a linear increase in CH4 fluxes with decreasing hydrostatic pressure, other authors (e.g., Iwata et al. 2020; Deshmukh et al. 2014) have suggested that CH4 bubbling sites are seasonally controlled by hydrostatic pressure. In Cubillas, CH4 fluxes are primarily seasonally controlled by temperature; however, during periods of declining water level, a linear relationship is still observed between CH4 fluxes and hydrostatic pressure (Figure S17). Similarly, temperature and depth continue to modulate CH4 emissions at the multiday scale, consistent with observations in other systems (Hounshell et al. 2023; Knox et al. 2021).
The diurnal variability of CH4 fluxes in Cubillas is better predicted by wind speed, near‐sediment temperature, and the temporal gradient of total pressure (ΔP tot/Δt). Wind‐driven turbulence enhances gas transfer velocity at the air–water interface, controlling diffusive CH4 fluxes (e.g., Lorke and Peeters 2006; Crusius and Wanninkhof 2003). Accordingly, CH4 diffusive fluxes in Cubillas vary synchronously with wind forcing at the diurnal scale (Figure 5c; León‐Palmero et al. 2025). Surface cooling can generate convective turbulence that also affects gas exchange (e.g., Tedford et al. 2014; Read et al. 2012; Soloviev et al. 2007), although RF analysis indicates that, on an annual basis, buoyancy‐driven convection plays a secondary role overall in Cubillas, in contrast to other systems (e.g., Podgrajsek et al. 2014). Wind can also induce bottom currents that enhance near‐bed shear stress, thereby favoring the release of gas bubbles from the sediment (Joyce and Jewell 2003). Temporal decreases in total pressure are well‐known to control ebullition (e.g., Varadharajan and Hemond 2012; Mattson and Likens 1990), which in Cubillas exhibits a clear diurnal bimodal pattern consistent with semidiurnal atmospheric pressure oscillations (Figure 5b), and similar to that reported by Deshmukh et al. (2014) in a tropical hydroelectric reservoir. Near‐sediment temperatures, pressure drops, and wind speed tend to peak during daytime hours, contributing to higher daytime CH4 fluxes (by ~ 50% or ~ 20%, depending on the year; Table 1; Figure S16), consistent with daytime‐elevated methane emissions reported for four lakes by Sieczko et al. (2020), where higher daytime wind speeds were identified as the dominant driver.
Average annual CO2 emissions in Cubillas fall within the wide range reported for temperate (101–103 mgC m−2 d−1 in absolute values, e.g., Prairie et al. 2021) and Mediterranean (León‐Palmero, Morales‐Baquero, and Reche 2020) reservoirs, with diurnal variability dominating total flux variance. CO2 fluxes in Cubillas fall within Cluster A identified by de Eyto et al. (2025) from 41 diel campaigns across 21 lakes, characterized by afternoon maxima and dawn minima, and primarily driven by wind forcing. The predominance of wind forcing as a driver for CO2 fluxes is also consistent with previous flux‐chamber measurements in the study reservoir (León‐Palmero et al. 2025; Rodríguez‐Velasco et al. 2024) and other year‐long EC studies in lakes and reservoirs (e.g., Golub et al. 2023). Multiday CO2 flux variability was primarily associated with water depth, temperature, chlorophyll‐a concentration, and GPP, highlighting the combined influence of hydrological conditions and ecosystem metabolism on CO2 dynamics. Air and water temperatures are common predictors of CO2 fluxes across multiple temporal scales in lakes and reservoirs (Rodríguez‐Velasco et al. 2024; Golub et al. 2023; Reed et al. 2018; Ouyang et al. 2017). Temperature likely controls short‐term respiration and organic carbon mineralization rates, while depth affects how CO2 and dissolved inorganic carbon are distributed and buffered within the water column. Chlorophyll‐a, as a proxy for phytoplankton biomass, influences CO2 fluxes through photosynthetic uptake, respiration, and the supply of labile organic carbon for microbial processing. The influence of chlorophyll‐a concentration at multiday timescales is consistent with previous studies; for example, Ouyang et al. (2017) and Shao et al. (2015) reported negative correlations between monthly CO2 fluxes and chlorophyll‐a concentrations in a western bay of Lake Erie. The emergence of GPP as the second major predictor of multiday variability in the wet year aligns with previous findings in aquatic systems, where air–water CO2 fluxes track short‐term variations in primary production rather than standing biomass (e.g., Wang et al. 2026).
Inflow discharge was not a significant predictor of seasonal CO2 flux variability, suggesting that short‐term external carbon inputs are not the dominant control on seasonal emissions. Instead, water depth—the major predictor of the seasonal component of the CO2 flux—likely integrates the cumulative effects of hydrology and carbon storage through pelagic respiration, while surface water temperature, the second most important predictor, regulates internal metabolic processing and CO2 availability. During the wet year, the reservoir rewetted areas that had been dry for more than 1 year, as maximum water levels declined from 2021 to 2023 (Figure 1a). For example, 5% of the in‐reservoir 90% EC footprint during the wet year corresponded to areas that were rewetted above the maximum water level reached in the previous year. Rewetting of previously dry areas may have promoted higher dissolved CO2 concentrations within the EC footprint, either through the presence of rewetted areas within the footprint itself or through enhanced advection of CO2‐rich water from other rewetted reservoir locations. These processes likely contributed to the early‐season peak in CO2 fluxes observed during the wet year by increasing substrate availability and microbial activation in already warm (T surf > 16°C) rewetted areas (the “Birch effect”, Amaral et al. 2022; Jarvis et al. 2007; Birch 1958). For example, rain events have been shown to cause at least a 5‐fold increase in CO2 fluxes compared to pre‐rainfall values in boreal forests (Makhnykina et al. 2024). Hysteresis between CO2 fluxes and surface water temperature during the wet year may have been further reinforced by the rapid summer decline in water level (4.5 m from June to September), leading to the flushing of water, nutrients, and carbon. Consequently, CO2 fluxes exhibited hysteresis with respect to surface water temperature, with higher emissions at a given temperature during the rising (rewetting) phase than during the falling (drawdown) phase, reflecting temporal changes in carbon availability rather than temperature alone. Additional measurements are needed to confirm this mechanism.
Although RF analysis does not explicitly account for dependencies among drivers or assign causation, we are confident that our conclusions regarding the key role of hydrological and thermal variability controlling GHG emissions from shallow eutrophic Mediterranean reservoirs are robust and consistent with our mechanistic understanding of GHG dynamics. Interannual differences in CH4 emissions were primarily driven by changes in water level and hydrostatic pressure, which regulated the timing and intensity of the high‐emission season via effects on ebullition and, in the case of water level, thermal stratification and sediment temperature. Lower water column depth and an earlier drawdown during the dry year led to substantially higher cumulative methane emissions (44% higher), highlighting how projected increases in drought frequency and altered hydrological regimes may enhance reservoir CH4 emissions. Water level dynamics are also influenced by reservoir operation, so management strategies may partly modulate these responses, although reduced summer withdrawals during drought may be unfeasible in water‐limited Mediterranean regions. Drought‐induced changes in reservoir surface area must also be considered when scaling GHG emissions at the reservoir level. In contrast to methane, CO2 fluxes were less sensitive to interannual hydrological variability and were dominated by short‐term physical drivers, particularly wind speed, although seasonal water‐level changes still influenced CO2 emissions, emphasizing how water‐column depth affects carbon storage and subsequent CO2 release. Finally, chlorophyll‐a–linked seasonal and multiday variability in CH4 and CO2 fluxes indicates that eutrophication enhances GHG emissions by boosting primary production and labile sedimentary carbon supply, reinforcing methane hysteresis and ebullitive temperature sensitivity.
Mediterranean‐climate regions are among the most densely populated and water‐stressed globally, relying on reservoirs to buffer seasonal and interannual water scarcity. The Mediterranean region is also a climate change hotspot, warming at rates that exceed global averages (e.g., Hassoun et al. 2025; Pausas and Millán 2019), but similar highly variable hydrological conditions are increasingly observed in other regions, with substantial portions of the global land surface transitioning toward drier aridity classes (Crapart et al. 2026). Our findings show that the mechanisms driving the observed interannual variability in CH4 emissions are broadly applicable to managed freshwater systems and relevant to regions experiencing increasing aridity. Importantly, we observed a clear decoupling in GHG dynamics, whereby drought conditions disproportionately enhanced emissions of the high‐radiative‐forcing gas CH4 while having little effect on CO2 fluxes, thereby substantially amplifying the warming impact per unit of stored water. By relying on averaged or static hydrology, global upscaling approaches may miss interannual variability, adding uncertainty to estimates of freshwater–climate feedbacks. Although the limited range of hydrological conditions examined here precludes establishing robust quantitative relationships, our findings indicate that future changes in reservoir GHG emissions will depend on the combined effects of climate‐driven and anthropogenic‐driven hydrological changes alongside increasing nutrient enrichment. This underscores the need for continued long‐term, high‐frequency measurements across a broader range of hydrological conditions to improve emission estimates and projections of climate feedbacks from managed freshwater systems.
Author Contributions
Cintia L. Ramón: conceptualization, data curation, formal analysis, funding acquisition, investigation, methodology, project administration, resources, software, validation, visualization, writing – original draft. Isabel Reche: conceptualization, funding acquisition, investigation, project administration, resources, supervision, writing – original draft, writing – review and editing. Rodrigo J. Gonçalves: data curation, investigation, software, writing – review and editing. Andrés Martínez‐García: data curation, investigation, writing – review and editing. Miriam García‐Alguacil: data curation, investigation, writing – review and editing. Sergio López‐Padilla: data curation, investigation, writing – review and editing. Javier Herrero: data curation, investigation, writing – review and editing. Rafael Morales‐Baquero: investigation, writing – review and editing. Enrique P. Sánchez‐Cañete: methodology, software. Alicia Cortés: investigation, writing – review and editing. Sara Valiente: data curation, investigation, writing – review and editing. Francisco J. Rueda: conceptualization, funding acquisition, investigation, project administration, resources, writing – review and editing.
Conflicts of Interest
The authors declare no conflicts of interest.
Supporting information
Figure S1: Annual footprints of the eddy covariance fluxes. Areas contributing to different percentages of the EC footprint centered on the EC tower location (colored lines), the bathymetry of the Cubillas Reservoir expressed as isolevels of water elevation (m.a.s.l.; gray color scale), and wind roses for (a) 2022 (dry year) and (b) 2024 (wet year). Thick and thin black contour lines indicate the minimum and maximum water levels reached in each year, respectively. Map lines delineate study areas and do not necessarily depict accepted national boundaries.
Figure S2: Rotated vs non‐rotated CH4 and CO2 fluxes. Comparison of 30‐min fluxes calculated with and without accounting for pitch and roll rotation of the floating platform. The black solid line indicates the 1:1 relationship. Root mean square errors are shown at the top of each panel. Differences between rotated and non‐rotated fluxes were tested using a linear mixed‐effects model, with rotation included as a fixed effect and day as a random intercept. No significant differences were detected between the two signals (p > 0.05).
Figure S3: Gap‐filled hourly time series of total CH4 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S4: Gap‐filled hourly time series of ebullitive CH4 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S5: Gap‐filled hourly time series of diffusive CH4 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S6: Gap‐filled hourly time series of CO2 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S7: Variables used as predictors in the random forest analysis. Variables: T air = Air temperature (°C). T surf = Surface temperature (°C), 0.5 m below the surface. T bot = Bottom temperature (°C), 0.5 m above the sediment. ΔT = Stratification strength (°C) (=T surf−T bot). P atm = Atmospheric pressure (hPa). Depth = Water column depth (m). ΔP tot/Δt = Gradient of total pressure (atmospheric + hydrostatic) (Pa h−1). Wind = Wind speed (m s−1). O 2,bot = Oxygen concentration 0.5 m above the sediment (mg L−1). Chl‐a = Chlorophyll‐a (μg L−1). NEP = Net ecosystem production (gO2 m−3 d−1). GPP = Gross primary production (gO2 m−3 d−1). Inflow = Inflow discharge (m3 s−1). B 0,net = Net surface buoyancy flux (W kg−1).
Figure S8: Spearman's correlation coefficients for predictor variables for Year 2022 (dry year). Statistical significance: *p < 0.05; **p < 0.01; ***p < 0.001. Variables: T air = Air temperature. T surf = Surface temperature (°C), 0.5 m below the surface. T bot = Bottom temperature (°C), 0.5 m above the sediment. ΔT = Stratification strength (°C) (= T surf−T bot). P atm = Atmospheric pressure (Pa). Depth = Water column depth (m). ΔP tot/Δt = Gradient of total pressure (atmospheric + hydrostatic) (Pa h−1). Wind = Wind speed (m s−1). O 2,bot = Oxygen concentration 0.5 m above the sediment (mg L−1). Chl‐a = Chlorophyll‐a (μg L−1). NEP = Net ecosystem production (gO2 m−3 d−1). GPP = Gross primary production (gO2 m−3 d−1). Inflow = Inflow discharge (m3 s−1). B 0,net = Net surface buoyancy flux (W kg−1). SWR = shortwave radiation (W m−2).
Figure S9: Reservoir‐wide total emissions. (a, b) Reservoir‐wide cumulative (a) CH4 and (b) CO2 emissions, expressed in moles and assuming that the EC footprint is representative of emissions from the entire reservoir, and (c) reservoir surface area during the wet and dry years.
Figure S10: Random forest predictions for the original and decomposed CH4 flux signals during the dry year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S11: Random Forest predictions for the original and decomposed CH4 flux signals during the wet year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S12: Random Forest predictions for the original and decomposed CO2 flux signals during the dry year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S13: Random forest predictions for the original and decomposed CO2 flux signals during the wet year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S14: Bottom water temperature vs. methane ebullitive and diffusive fluxes. The black dashed line shows the best 2‐year Arrhenius fit [Equation (1)], with the corresponding activation energy (E A ) indicated. The gray shaded area around each Arrhenius fit represents the 95% confidence interval.
Figure S15: Random forest predictor importance for the decomposed diurnal diffusive (a) and ebullitive (b) CH4 fluxes. Diffusive and ebullitive components were extracted from the total eddy covariance CH4 fluxes using the wavelet analysis approach proposed by Iwata et al. (2018). The decomposed temporal flux components were obtained using the maximal overlap discrete wavelet transform (MODWT) and multiresolution analysis. Each Random Forest analysis was repeated 50 times using different random seeds to assess model stochasticity. Bars represent mean predictor importance across runs, and black horizontal lines indicate ±1 standard deviation.
Figure S16: Daytime versus nighttime fluxes. Daily‐averaged gap‐filled CH4 (a, b) and CO2 (c, d) fluxes during daytime (PAR ≥ 10 μmol photons m−2 s−1) and nighttime (PAR < 10 μmol photons m−2 s−1) conditions for the dry year (a, c) and the wet year (b, d).
Figure S17: Hydrostatic pressure versus methane fluxes. Relationship between daily‐averaged hydrostatic pressure at the lake sediment surface and methane fluxes. Solid gray lines show the seasonally decomposed relationship between the two variables, and arrows indicate the direction of progression over the annual cycle. Linear regressions are shown for periods during which CH4 fluxes increase linearly with decreasing hydrostatic pressure (i.e., water depth). The gray shaded area around each regression represents the 95% confidence interval.
Table S1: Coefficient of determination (R 2) for random forest predictions as the number of variables (within brackets) increases from 1 to 5, or until a combination reaching R 2 > 0.95 is achieved. Variables 1–5 were selected based on their predicted importance (the most important first).
Acknowledgements
Grant PID2022‐137865OB‐I00—funded by MICIU/AEI/10.13039/501100011033 and by ERDF/EU—, project COSTUMER (ref. P21_00238), —funded by the Regional Government of Andalusia (Junta de Andalucía, Consejería de Universidad, and Investigación e Innovación)—, and Grant TED2021‐130744B‐C22—funded by MICIU/AEI/10.13039/501100011033 and by the European Union Next Generation EU/PRTR—supported this study. Funding for open access charge: Universidad de Granada.
Contributor Information
Cintia L. Ramón, Email: crcasanas@ugr.es.
Isabel Reche, Email: ireche@ugr.es.
Data Availability Statement
The data supporting the findings of this study are openly available in https://doi.org/10.5281/zenodo.20800380.
References
- Aben, R. C. H. , Barros N., Van Donk E., et al. 2017. “Cross Continental Increase in Methane Ebullition Under Climate Change.” Nature Communications 8, no. 1: 1–8. 10.1038/s41467-017-01535-y. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Amaral, J. H. F. , Melack J. M., Barbosa P. M., et al. 2022. “Inundation, Hydrodynamics and Vegetation Influence Carbon Dioxide Concentrations in Amazon Floodplain Lakes.” Ecosystems 25, no. 4: 911–930. 10.1007/S10021-021-00692-Y. [DOI] [Google Scholar]
- Baldocchi, D. 2014. “Measuring Fluxes of Trace Gases and Energy Between Ecosystems and the Atmosphere – The State and Future of the Eddy Covariance Method.” Global Change Biology 20, no. 12: 3600–3609. 10.1111/GCB.12649. [DOI] [PubMed] [Google Scholar]
- Bastviken, D. , Cole J., Pace M., and Tranvik L.. 2004. “Methane Emissions From Lakes: Dependence of Lake Characteristics, Two Regional Assessments, and a Global Estimate.” Global Biogeochemical Cycles 18, no. 4: 1–12. 10.1029/2004GB002238. [DOI] [Google Scholar]
- Birch, H. F. 1958. “The Effect of Soil Drying on Humus Decomposition and Nitrogen Availability.” Plant and Soil 10, no. 1: 9–31. 10.1007/BF01343734. [DOI] [Google Scholar]
- Chang, K. Y. , Riley W. J., Crill P. M., Grant R. F., and Saleska S. R.. 2020. “Hysteretic Temperature Sensitivity of Wetland CH4 Fluxes Explained by Substrate Availability and Microbial Activity.” Biogeosciences 17, no. 22: 5849–5860. 10.5194/BG-17-5849-2020. [DOI] [Google Scholar]
- Chang, K. Y. , Riley W. J., Knox S. H., et al. 2021. “Substantial Hysteresis in Emergent Temperature Sensitivity of Global Wetland CH4 Emissions.” Nature Communications 12, no. 1: 1–10. 10.1038/s41467-021-22452-1. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Cole, J. J. , Prairie Y. T., Caraco N. F., et al. 2007. “Plumbing the Global Carbon Cycle: Integrating Inland Waters Into the Terrestrial Carbon Budget.” Ecosystems 10, no. 1: 171–184. 10.1007/S10021-006-9013-8. [DOI] [Google Scholar]
- Crapart, C. , Anquetin S., Blanchet J., and Diedhiou A.. 2026. “Global Projections of Aridity Index for Mid and Long‐Term Future Based on CMIP6 Scenarios.” Hydrology and Earth System Sciences 30, no. 1: 163–181. 10.5194/HESS-30-163-2026. [DOI] [Google Scholar]
- Crusius, J. , and Wanninkhof R.. 2003. “Gas Transfer Velocities Measured at Low Wind Speed Over a Lake.” Limnology and Oceanography 48, no. 3: 1010–1017. 10.4319/LO.2003.48.3.1010. [DOI] [Google Scholar]
- de Eyto, E. , Smyth R. L., Pilla R. M., et al. 2025. “Diel Variation in CO2 Flux Is Substantial in Many Lakes.” Limnology and Oceanography Letters 10, no. 6: 977–989. 10.1002/lol2.70066. [DOI] [Google Scholar]
- Deemer, B. R. , Harrison J. A., Li S., et al. 2016. “Greenhouse Gas Emissions From Reservoir Water Surfaces: A New Global Synthesis.” Bioscience 66, no. 11: 949–964. 10.1093/biosci/biw117. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Deemer, B. R. , and Holgerson M. A.. 2021. “Drivers of Methane Flux Differ Between Lakes and Reservoirs, Complicating Global Upscaling Efforts.” Journal of Geophysical Research: Biogeosciences 126, no. 4: e2019JG005600. 10.1029/2019JG005600. [DOI] [Google Scholar]
- DelSontro, T. , Beaulieu J. J., and Downing J. A.. 2018. “Greenhouse Gas Emissions From Lakes and Impoundments: Upscaling in the Face of Global Change.” Limnology and Oceanography Letters 3, no. 3: 64–75. 10.1002/lol2.10073. [DOI] [PMC free article] [PubMed] [Google Scholar]
- DelSontro, T. , Boutet L., St‐Pierre A., del Giorgio P. A., and Prairie Y. T.. 2016. “Methane Ebullition and Diffusion From Northern Ponds and Lakes Regulated by the Interaction Between Temperature and System Productivity.” Limnology and Oceanography 61, no. S1: S62–S77. 10.1002/LNO.10335. [DOI] [Google Scholar]
- Deshmukh, C. , Serça D., Delon C., et al. 2014. “Physical Controls on CH4 Emissions From a Newly Flooded Subtropical Freshwater Hydroelectric Reservoir: Nam Theun 2.” Biogeosciences 11, no. 15: 4251–4269. 10.5194/BG-11-4251-2014. [DOI] [Google Scholar]
- Erkkilä, K. M. , Ojala A., Bastviken D., et al. 2018. “Methane and Carbon Dioxide Fluxes Over a Lake: Comparison Between Eddy Covariance, Floating Chambers and Boundary Layer Method.” Biogeosciences 15, no. 2: 429–445. 10.5194/BG-15-429-2018. [DOI] [Google Scholar]
- Esteban‐Parra, M. J. , García‐Valdecasas Ojeda M., Peinó‐Calero E., et al. 2022. “Climate Variability and Trends.” In The Landscape of the Sierra Nevada: A Unique Laboratory of Global Processes in Spain, 129–148. Springer. 10.1007/978-3-030-94219-9_9. [DOI] [Google Scholar]
- Eugster, W. , Delsontro T., Shaver G. R., and Kling G. W.. 2020. “Interannual, Summer, and Diel Variability of CH4 and CO2 Effluxes From Toolik Lake, Alaska, During the Ice‐Free Periods 2010–2015.” Environmental Science: Processes & Impacts 22, no. 11: 2181–2198. 10.1039/D0EM00125B. [DOI] [PubMed] [Google Scholar]
- Eugster, W. , Kling G., Jonas T., et al. 2003. “CO2 Exchange Between Air and Water in an Arctic Alaskan and Midlatitude Swiss Lake: Importance of Convective Mixing.” Journal of Geophysical Research: Atmospheres 108, no. D12: 4362. 10.1029/2002JD002653. [DOI] [Google Scholar]
- Golub, M. , Koupaei‐Abyazani N., Vesala T., et al. 2023. “Diel, Seasonal, and Inter‐Annual Variation in Carbon Dioxide Effluxes From Lakes and Reservoirs.” Environmental Research Letters 18, no. 3: 34046. 10.1088/1748-9326/ACB834. [DOI] [Google Scholar]
- Grasset, C. , Mendonça R., Villamor Saucedo G., Bastviken D., Roland F., and Sobek S.. 2018. “Large but Variable Methane Production in Anoxic Freshwater Sediment Upon Addition of Allochthonous and Autochthonous Organic Matter.” Limnology and Oceanography 63, no. 4: 1488–1501. 10.1002/LNO.10786. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Hassoun, A. E. R. , Mojtahid M., Merheb M., Lionello P., Gattuso J. P., and Cramer W.. 2025. “Climate Change Risks on Key Open Marine and Coastal Mediterranean Ecosystems.” Scientific Reports 15, no. 1: 24907. 10.1038/s41598-025-07858-x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Herrero, J. 2007. “Modelo físico de acumulación y fusión de la nieve.” In Aplicación en Sierra Nevada (España). University of Granada. http://hdl.handle.net/10481/1729. [Google Scholar]
- Hounshell, A. G. , D'Acunha B. M., Breef‐Pilz A., Johnson M. S., Thomas R. Q., and Carey C. C.. 2023. “Eddy Covariance Data Reveal That a Small Freshwater Reservoir Emits a Substantial Amount of Carbon Dioxide and Methane.” Journal of Geophysical Research: Biogeosciences 128, no. 3: e2022JG007091. 10.1029/2022JG007091. [DOI] [Google Scholar]
- Huotari, J. , Ojala A., Peltomaa E., et al. 2011. “Long‐Term Direct CO2 Flux Measurements Over a Boreal Lake: Five Years of Eddy Covariance Data.” Geophysical Research Letters 38, no. 18: L18401. 10.1029/2011GL048753. [DOI] [Google Scholar]
- Iwata, H. , Hirata R., Takahashi Y., Miyabara Y., Itoh M., and Iizuka K.. 2018. “Partitioning Eddy‐Covariance Methane Fluxes From a Shallow Lake Into Diffusive and Ebullitive Fluxes.” Boundary‐Layer Meteorology 169, no. 3: 413–428. 10.1007/s10546-018-0383-1. [DOI] [Google Scholar]
- Iwata, H. , Nakazawa K., Sato H., et al. 2020. “Temporal and Spatial Variations in Methane Emissions From the Littoral Zone of a Shallow Mid‐Latitude Lake With Steady Methane Bubble Emission Areas.” Agricultural and Forest Meteorology 295: 108184. 10.1016/J.AGRFORMET.2020.108184. [DOI] [Google Scholar]
- Jammet, M. , Crill P., Dengel S., and Friborg T.. 2015. “Large Methane Emissions From a Subarctic Lake During Spring Thaw: Mechanisms and Landscape Significance.” Journal of Geophysical Research: Biogeosciences 120, no. 11: 2289–2305. 10.1002/2015JG003137. [DOI] [Google Scholar]
- Jammet, M. , Dengel S., Kettner E., et al. 2017. “Year‐Round CH4 and CO2 Flux Dynamics in Two Contrasting Freshwater Ecosystems of the Subarctic.” Biogeosciences 14, no. 22: 5189–5216. 10.5194/BG-14-5189-2017. [DOI] [Google Scholar]
- Jansen, J. , Thornton B. F., Wik M., MacIntyre S., and Crill P. M.. 2020. “Temperature Proxies as a Solution to Biased Sampling of Lake Methane Emissions.” Geophysical Research Letters 47, no. 14: e2020GL088647. 10.1029/2020GL088647. [DOI] [Google Scholar]
- Jarvis, P. , Rey A., Petsikos C., et al. 2007. “Drying and Wetting of Mediterranean Soils Stimulates Decomposition and Carbon Dioxide Emission: The “Birch Effect”.” Tree Physiology 27, no. 7: 929–940. 10.1093/TREEPHYS/27.7.929. [DOI] [PubMed] [Google Scholar]
- Johnson, M. S. , Matthews E., Bastviken D., Deemer B., Du J., and Genovese V.. 2021. “Spatiotemporal Methane Emission From Global Reservoirs.” Journal of Geophysical Research: Biogeosciences 126, no. 8: e2021JG006305. 10.1029/2021JG006305. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Johnson, M. S. , Matthews E., Du J., Genovese V., and Bastviken D.. 2022. “Methane Emission From Global Lakes: New Spatiotemporal Data and Observation‐Driven Modeling of Methane Dynamics Indicates Lower Emissions.” Journal of Geophysical Research: Biogeosciences 127, no. 7: e2022JG006793. 10.1029/2022JG006793. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Joyce, J. , and Jewell P. W.. 2003. “Physical Controls on Methane Ebullition From Reservoirs and Lakes.” Environmental and Engineering Geoscience 9, no. 2: 167–178. 10.2113/9.2.167. [DOI] [Google Scholar]
- Kim, B.‐K. , Kim H., Won H., et al. 2025. Effects of Air Temperature, Atmospheric Pressure, and Wind Speed on Methane Emissions in Yonghwasil Reservoir. Environmental Science & Technology Letters. 10.1021/ACS.ESTLETT.5C00535. [DOI] [Google Scholar]
- Kljun, N. , Calanca P., Rotach M. W., and Schmid H. P.. 2015. “A Simple Two‐Dimensional Parameterisation for Flux Footprint Prediction (FFP).” Geoscientific Model Development 8, no. 11: 3695–3713. 10.5194/GMD-8-3695-2015. [DOI] [Google Scholar]
- Knox, S. H. , Bansal S., McNicol G., et al. 2021. “Identifying Dominant Environmental Predictors of Freshwater Wetland Methane Fluxes Across Diurnal to Seasonal Time Scales.” Global Change Biology 27, no. 15: 3582–3604. 10.1111/GCB.15661. [DOI] [PubMed] [Google Scholar]
- Knox, S. H. , Jackson R. B., Poulter B., et al. 2019. “FLUXNET‐CH4 Synthesis Activity: Objectives, Observations, and Future Directions.” Bulletin of the American Meteorological Society 100, no. 12: 2607–2632. 10.1175/BAMS-D-18-0268.1. [DOI] [Google Scholar]
- Lapierre, J. F. , Guillemette F., Berggren M., and Del Giorgio P. A.. 2013. “Increases in Terrestrially Derived Carbon Stimulate Organic Carbon Processing and CO2 Emissions in Boreal Aquatic Ecosystems.” Nature Communications 4, no. 1: 1–7. 10.1038/ncomms3972. [DOI] [PubMed] [Google Scholar]
- Lehner, B. , and Döll P.. 2004. “Development and Validation of a Global Database of Lakes, Reservoirs and Wetlands.” Journal of Hydrology 296, no. 1–4: 1–22. 10.1016/J.JHYDROL.2004.03.028. [DOI] [Google Scholar]
- León‐Palmero, E. , Contreras‐Ruiz A., Sierra A., Morales‐Baquero R., and Reche I.. 2020. “Dissolved CH4 Coupled to Photosynthetic Picoeukaryotes in Oxic Waters and to Cumulative Chlorophyll‐a in Anoxic Waters of Reservoirs.” Biogeosciences 17, no. 12: 3223–3245. 10.5194/bg-17-3223-2020. [DOI] [Google Scholar]
- León‐Palmero, E. , Morales‐Baquero R., and Reche I.. 2020. “Greenhouse Gas Fluxes From Reservoirs Determined by Watershed Lithology, Morphometry, and Anthropogenic Pressure.” Environmental Research Letters 15, no. 4: 44012. 10.1088/1748-9326/ab7467. [DOI] [Google Scholar]
- León‐Palmero, E. , Morales‐Baquero R., and Reche I.. 2025. “Higher Emissions of Carbon Dioxide, Nitrous Oxide, and Methane During the Daytime in Two Reservoirs.” Biogeochemistry 168, no. 6: 99. 10.1007/S10533-025-01283-Y. [DOI] [Google Scholar]
- Liu, H. , Zhang Q., Katul G. G., Cole J. J., Chapin F. S., and MacIntyre S.. 2016. “Large CO2 Effluxes at Night and During Synoptic Weather Events Significantly Contribute to CO2 Emissions From a Reservoir.” Environmental Research Letters 11, no. 6: 64001. 10.1088/1748-9326/11/6/064001. [DOI] [Google Scholar]
- Lorke, A. , and Peeters F.. 2006. “Toward a Unified Scaling Relation for Interfacial Fluxes.” Journal of Physical Oceanography 36, no. 5: 955–961. 10.1175/JPO2903.1. [DOI] [Google Scholar]
- MacIntyre, S. , Jonsson A., Jansson M., Aberg J., Turney D. E., and Miller S. D.. 2010. “Buoyancy Flux, Turbulence, and the Gas Transfer Coefficient in a Stratified Lake.” Geophysical Research Letters 37, no. 24: 24604. 10.1029/2010GL044164. [DOI] [Google Scholar]
- Makhnykina, A. V. , Vaganov E. A., Panov A. V., Koshurnikova N. N., and Prokushkin A. S.. 2024. “The Pulses of Soil CO2 Emission in Response to Rainfall Events in Central Siberia: Revisiting the Overall Frost‐Free Season CO2 Flux.” Forests 15, no. 2: 355. 10.3390/F15020355. [DOI] [Google Scholar]
- Mammarella, I. , Nordbo A., Rannik Ü., et al. 2015. “Carbon Dioxide and Energy Fluxes Over a Small Boreal Lake in Southern Finland.” Journal of Geophysical Research: Biogeosciences 120, no. 7: 1296–1314. 10.1002/2014JG002873. [DOI] [Google Scholar]
- Many, G. , Escoffier N., Perolo P., Bärenbold F., Bouffard D., and Perga M.‐E.. 2024. “Calcite Precipitation: The Forgotten Piece of Lakes' Carbon Cycle.” Science Advances 10, no. 44: eado5924. 10.1126/SCIADV.ADO5924. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Martínez‐García, A. , Peralta‐Maraver I., Rodríguez‐Velasco E., et al. 2024. “Particulate Organic Carbon Sedimentation Triggers Lagged Methane Emissions in a Eutrophic Reservoir.” Limnology and Oceanography Letters 9, no. 3: 247–257. 10.1002/LOL2.10379. [DOI] [Google Scholar]
- Mattson, M. D. , and Likens G. E.. 1990. “Air Pressure and Methane Fluxes.” Nature 347, no. 6295: 718–719. 10.1038/347718b0. [DOI] [Google Scholar]
- Mauder, M. , and Foken T.. 2004. “Documentation and Instruction Manual of the Eddy Covariance Software Package TK2.” Arbeit, 26.
- Mayr, M. J. , Zimmermann M., Dey J., Brand A., Wehrli B., and Bürgmann H.. 2020. “Growth and Rapid Succession of Methanotrophs Effectively Limit Methane Release During Lake Overturn.” Communications Biology 3, no. 1: 108. 10.1038/s42003-020-0838-z. [DOI] [PMC free article] [PubMed] [Google Scholar]
- McDonald, C. P. , Stets E. G., Striegl R. G., and Butman D.. 2013. “Inorganic Carbon Loading as a Primary Driver of Dissolved Carbon Dioxide Concentrations in the Lakes and Reservoirs of the Contiguous United States.” Global Biogeochemical Cycles 27, no. 2: 285–295. 10.1002/GBC.20032. [DOI] [Google Scholar]
- Messager, M. L. , Lehner B., Grill G., Nedeva I., and Schmitt O.. 2016. “Estimating the Volume and Age of Water Stored in Global Lakes Using a Geo‐Statistical Approach.” Nature Communications 7, no. 1: 1–11. 10.1038/ncomms13603. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Moncrieff, J. , Clement R., Finnigan J., and Meyers T.. 2004. “Averaging, Detrending, and Filtering of Eddy Covariance Time Series.” In Handbook of Micrometeorology, 7–31. Springer. 10.1007/1-4020-2265-4_2. [DOI] [Google Scholar]
- Moncrieff, J. B. , Massheder J. M., De Bruin H., et al. 1997. “A System to Measure Surface Fluxes of Momentum, Sensible Heat, Water Vapour and Carbon Dioxide.” Journal of Hydrology 188–189, no. 1–4: 589–611. 10.1016/S0022-1694(96)03194-0. [DOI] [Google Scholar]
- Natchimuthu, S. , Sundgren I., Gålfalk M., et al. 2016. “Spatio‐Temporal Variability of Lake CH4 Fluxes and Its Influence on Annual Whole Lake Emission Estimates.” Limnology and Oceanography 61, no. S1: S13–S26. 10.1002/LNO.10222. [DOI] [Google Scholar]
- Niu, X. , Yan Z., Wu W., Hua Z., Hao L., and Rosentreter J. A.. 2025. “Rainfall Can Significantly Reduce Pond Methane Emissions by Depressing Ebullition.” Water Research 285: 124167. 10.1016/J.WATRES.2025.124167. [DOI] [PubMed] [Google Scholar]
- Ordóñez, C. , DelSontro T., Langenegger T., Donis D., Suarez E. L., and McGinnis D. F.. 2023. “Evaluation of the Methane Paradox in Four Adjacent Pre‐Alpine Lakes Across a Trophic Gradient.” Nature Communications 14, no. 1: 1–13. 10.1038/s41467-023-37861-7. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Ouyang, Z. , Shao C., Chu H., et al. 2017. “The Effect of Algal Blooms on Carbon Emissions in Western Lake Erie: An Integration of Remote Sensing and Eddy Covariance Measurements.” Remote Sensing 9, no. 1: 44. 10.3390/RS9010044. [DOI] [Google Scholar]
- Pausas, J. G. , and Millán M. M.. 2019. “Greening and Browning in a Climate Change Hotspot: The Mediterranean Basin.” BioScience 69, no. 2: 143–151. 10.1093/BIOSCI/BIY157. [DOI] [Google Scholar]
- Podgrajsek, E. , Sahlée E., Bastviken D., et al. 2016. “Methane Fluxes From a Small Boreal Lake Measured With the Eddy Covariance Method.” Limnology and Oceanography 61, no. S1: S41–S50. 10.1002/LNO.10245. [DOI] [Google Scholar]
- Podgrajsek, E. , Sahlée E., and Rutgersson A.. 2014. “Diurnal Cycle of Lake Methane Flux.” Journal of Geophysical Research: Biogeosciences 119, no. 3: 236–248. 10.1002/2013JG002327. [DOI] [Google Scholar]
- Praetzel, L. S. E. , Schmiedeskamp M., and Knorr K. H.. 2021. “Temperature and Sediment Properties Drive Spatiotemporal Variability of Methane Ebullition in a Small and Shallow Temperate Lake.” Limnology and Oceanography 66, no. 7: 2598–2610. 10.1002/LNO.11775. [DOI] [Google Scholar]
- Prairie, Y. T. , Mercier‐Blais S., Harrison J. A., et al. 2021. “G‐Res Tool Modelling Database.” Zenodo. 10.5281/ZENODO.4711132. [DOI]
- Raymond, P. A. , Hartmann J., Lauerwald R., et al. 2013. “Global Carbon Dioxide Emissions From Inland Waters.” Nature 503, no. 7476: 355–359. 10.1038/nature12760. [DOI] [PubMed] [Google Scholar]
- Read, J. S. , Hamilton D. P., Desai A. R., et al. 2012. “Lake‐Size Dependency of Wind Shear and Convection as Controls on Gas Exchange.” Geophysical Research Letters 39, no. 9: L09405. 10.1029/2012GL051886. [DOI] [Google Scholar]
- Reed, D. E. , Dugan H. A., Flannery A. L., and Desai A. R.. 2018. “Carbon Sink and Source Dynamics of a Eutrophic Deep Lake Using Multiple Flux Observations Over Multiple Years.” Limnology and Oceanography Letters 3, no. 3: 285–292. 10.1002/LOL2.10075. [DOI] [Google Scholar]
- Rodríguez‐Velasco, E. , Peralta‐Maraver I., Martínez‐García A., et al. 2024. “Idiosyncratic Phenology of Greenhouse Gas Emissions in a Mediterranean Reservoir.” Limnology and Oceanography Letters 9: 364–375. 10.1002/LOL2.10409. [DOI] [Google Scholar]
- Rosentreter, J. A. , Borges A. V., Deemer B. R., et al. 2021. “Half of Global Methane Emissions Come From Highly Variable Aquatic Ecosystem Sources.” Nature Geoscience 14, no. 4: 225–230. 10.1038/s41561-021-00715-2. [DOI] [Google Scholar]
- Rousso, B. Z. , Bertone E., Stewart R. A., Rinke K., and Hamilton D. P.. 2021. “Light‐Induced Fluorescence Quenching Leads to Errors in Sensor Measurements of Phytoplankton Chlorophyll and Phycocyanin.” Water Research 198: 117133. 10.1016/J.WATRES.2021.117133. [DOI] [PubMed] [Google Scholar]
- Scholz, K. , Ejarque E., Hammerle A., Kainz M., Schelker J., and Wohlfahrt G.. 2021. “Atmospheric CO2 Exchange of a Small Mountain Lake: Limitations of Eddy Covariance and Boundary Layer Modeling Methods in Complex Terrain.” Journal of Geophysical Research: Biogeosciences 126, no. 7: e2021JG006286. 10.1029/2021JG006286. [DOI] [Google Scholar]
- Shao, C. , Chen J., Stepien C. A., et al. 2015. “Diurnal to Annual Changes in Latent, Sensible Heat, and CO2 Fluxes Over a Laurentian Great Lake: A Case Study in Western Lake Erie.” Journal of Geophysical Research: Biogeosciences 120, no. 8: 1587–1604. 10.1002/2015JG003025. [DOI] [Google Scholar]
- Sieczko, A. K. , Thanh Duc N., Schenk J., et al. 2020. “Diel Variability of Methane Emissions From Lakes.” Proceedings of the National Academy of Sciences of the United States of America 117, no. 35: 21488–21494. 10.1073/PNAS.2006024117. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Soloviev, A. , Donelan M., Graber H., Haus B., and Schlüssel P.. 2007. “An Approach to Estimation of Near‐Surface Turbulence and CO2 Transfer Velocity From Remote Sensing Data.” Journal of Marine Systems 66, no. 194: 182. 10.1016/J.JMARSYS.2006.03.023. [DOI] [Google Scholar]
- Spank, U. , Bernhofer C., Mauder M., Keller P. S., and Koschorreck M.. 2023. “Contrasting Temporal Dynamics of Methane and Carbon Dioxide Emissions From a Eutrophic Reservoir Detected by Eddy Covariance Measurements.” Meteorologische Zeitschrift 32, no. 4: 317–342. 10.1127/METZ/2023/1162. [DOI] [Google Scholar]
- Staehr, P. A. , Bade D., van de Bogert M. C., et al. 2010. “Lake Metabolism and the Diel Oxygen Technique: State of the Science.” Limnology and Oceanography: Methods 8, no. 11: 628–644. 10.4319/LOM.2010.8.0628. [DOI] [Google Scholar]
- Sturtevant, C. , Ruddell B. L., Knox S. H., et al. 2016. “Identifying Scale‐Emergent, Nonlinear, Asynchronous Processes of Wetland Methane Exchange.” Journal of Geophysical Research: Biogeosciences 121, no. 1: 188–204. 10.1002/2015JG003054. [DOI] [Google Scholar]
- Tang, K. W. , McGinnis D. F., Frindte K., Brüchert V., and Grossart H. P.. 2014. “Paradox Reconsidered: Methane Oversaturation in Well‐Oxygenated Lake Waters.” Limnology and Oceanography 59, no. 1: 275–284. 10.4319/LO.2014.59.1.0275. [DOI] [Google Scholar]
- Taoka, T. , Iwata H., Hirata R., Takahashi Y., Miyabara Y., and Itoh M.. 2020. “Environmental Controls of Diffusive and Ebullitive Methane Emissions at a Subdaily Time Scale in the Littoral Zone of a Midlatitude Shallow Lake.” Journal of Geophysical Research: Biogeosciences 125, no. 9: e2020JG005753. 10.1029/2020JG005753. [DOI] [Google Scholar]
- Tedford, E. W. , MacIntyre S., Miller S. D., and Czikowsky M. J.. 2014. “Similarity Scaling of Turbulence in a Temperate Lake During Fall Cooling.” Journal of Geophysical Research: Oceans 119, no. 8: 4689–4713. 10.1002/2014JC010135. [DOI] [Google Scholar]
- Tranvik, L. J. , Downing J. A., Cotner J. B., et al. 2009. “Lakes and Reservoirs as Regulators of Carbon Cycling and Climate.” Limnology and Oceanography 54, no. 6part2: 2298–2314. 10.4319/lo.2009.54.6_part_2.2298. [DOI] [Google Scholar]
- Updegraff, K. , Bridgham S. D., Pastor J., and Weishampel P.. 1998. “Hysteresis in the Temperature Response of Carbon Dioxide and Methane Production in Peat Soils.” Biogeochemistry 43, no. 3: 253–272. 10.1023/A:1006097808262. [DOI] [Google Scholar]
- Ustinov, N. B. , Agafonova S. A., and Kazantsev V. S.. 2025. “Methane Emissions From Lakes and Reservoirs: Uncertainties due to the Formation and Break‐Up of the Ice Cover.” Izvestiya, Atmospheric and Oceanic Physics 61, no. 4: 499–510. 10.1134/S0001433825700823. [DOI] [Google Scholar]
- Varadharajan, C. , and Hemond H. F.. 2012. “Time‐Series Analysis of High‐Resolution Ebullition Fluxes From a Stratified, Freshwater Lake.” Journal of Geophysical Research: Biogeosciences 117, no. G2: G02004. 10.1029/2011JG001866. [DOI] [Google Scholar]
- Vickers, D. , and Mahrt L.. 1997. “Quality Control and Flux Sampling Problems for Tower and Aircraft Data.” Journal of Atmospheric and Oceanic Technology 14, no. 3: 512–526. 10.1175/1520-0426(1997)014<0512:QCAFSP>2.0.CO;2. [DOI] [Google Scholar]
- Waldo, S. , Beaulieu J. J., Barnett W., et al. 2021. “Temporal Trends in Methane Emissions From a Small Eutrophic Reservoir: The Key Role of a Spring Burst.” Biogeosciences 18, no. 19: 5291–5311. 10.5194/BG-18-5291-2021. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Wang, Y. , Chen F., Chen C., et al. 2026. “Coupling Air–Water CO2 Flux and Primary Production Dynamics Under Hydrologic Variability in a Large Urban Estuary.” Limnology and Oceanography Letters 11, no. 1: e70072. 10.1002/LOL2.70072. [DOI] [Google Scholar]
- Webb, E. K. , Pearman G. I., and Leuning R.. 1980. “Correction of Flux Measurements for Density Effects due to Heat and Water Vapour Transfer.” Quarterly Journal of the Royal Meteorological Society 106, no. 447: 85–100. 10.1002/QJ.49710644707. [DOI] [Google Scholar]
- West, W. E. , Coloso J. J., and Jones S. E.. 2012. “Effects of Algal and Terrestrial Carbon on Methane Production Rates and Methanogen Community Structure in a Temperate Lake Sediment.” Freshwater Biology 57, no. 5: 949–955. 10.1111/J.1365-2427.2012.02755.X. [DOI] [Google Scholar]
- West, W. E. , Mccarthy S. M., and Jones S. E.. 2015. “Phytoplankton Lipid Content Influences Freshwater Lake Methanogenesis.” Freshwater Biology 60, no. 11: 2261–2269. 10.1111/FWB.12652. [DOI] [Google Scholar]
- Wik, M. , Crill P. M., Varner R. K., and Bastviken D.. 2013. “Multiyear Measurements of Ebullitive Methane Flux From Three Subarctic Lakes.” Journal of Geophysical Research: Biogeosciences 118, no. 3: 1307–1321. 10.1002/jgrg.20103. [DOI] [Google Scholar]
- Wik, M. , Thornton B. F., Bastviken D., Macintyre S., Varner R. K., and Crill P. M.. 2014. “Energy Input Is Primary Controller of Methane Bubbling in Subarctic Lakes.” Geophysical Research Letters 41, no. 2: 555–560. 10.1002/2013GL058510. [DOI] [Google Scholar]
- Wilczak, J. M. , Oncley S. P., and Stage S. A.. 2001. “Sonic Anemometer Tilt Correction Algorithms.” Boundary‐Layer Meteorology 99, no. 1: 127–150. 10.1023/A:1018966204465. [DOI] [Google Scholar]
- Wutzler, T. , Lucas‐Moffat A., Migliavacca M., et al. 2018. “Basic and Extensible Post‐Processing of Eddy Covariance Flux Data With REddyProc.” Biogeosciences 15, no. 16: 5015–5030. 10.5194/BG-15-5015-2018. [DOI] [Google Scholar]
- Yvon‐Durocher, G. , Allen A. P., Bastviken D., et al. 2014. “Methane Fluxes Show Consistent Temperature Dependence Across Microbial to Ecosystem Scales.” Nature 507, no. 7493: 488–491. 10.1038/nature13164. [DOI] [PubMed] [Google Scholar]
Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Figure S1: Annual footprints of the eddy covariance fluxes. Areas contributing to different percentages of the EC footprint centered on the EC tower location (colored lines), the bathymetry of the Cubillas Reservoir expressed as isolevels of water elevation (m.a.s.l.; gray color scale), and wind roses for (a) 2022 (dry year) and (b) 2024 (wet year). Thick and thin black contour lines indicate the minimum and maximum water levels reached in each year, respectively. Map lines delineate study areas and do not necessarily depict accepted national boundaries.
Figure S2: Rotated vs non‐rotated CH4 and CO2 fluxes. Comparison of 30‐min fluxes calculated with and without accounting for pitch and roll rotation of the floating platform. The black solid line indicates the 1:1 relationship. Root mean square errors are shown at the top of each panel. Differences between rotated and non‐rotated fluxes were tested using a linear mixed‐effects model, with rotation included as a fixed effect and day as a random intercept. No significant differences were detected between the two signals (p > 0.05).
Figure S3: Gap‐filled hourly time series of total CH4 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S4: Gap‐filled hourly time series of ebullitive CH4 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S5: Gap‐filled hourly time series of diffusive CH4 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S6: Gap‐filled hourly time series of CO2 fluxes and their decomposed seasonal, multiday, and diurnal components.
Figure S7: Variables used as predictors in the random forest analysis. Variables: T air = Air temperature (°C). T surf = Surface temperature (°C), 0.5 m below the surface. T bot = Bottom temperature (°C), 0.5 m above the sediment. ΔT = Stratification strength (°C) (=T surf−T bot). P atm = Atmospheric pressure (hPa). Depth = Water column depth (m). ΔP tot/Δt = Gradient of total pressure (atmospheric + hydrostatic) (Pa h−1). Wind = Wind speed (m s−1). O 2,bot = Oxygen concentration 0.5 m above the sediment (mg L−1). Chl‐a = Chlorophyll‐a (μg L−1). NEP = Net ecosystem production (gO2 m−3 d−1). GPP = Gross primary production (gO2 m−3 d−1). Inflow = Inflow discharge (m3 s−1). B 0,net = Net surface buoyancy flux (W kg−1).
Figure S8: Spearman's correlation coefficients for predictor variables for Year 2022 (dry year). Statistical significance: *p < 0.05; **p < 0.01; ***p < 0.001. Variables: T air = Air temperature. T surf = Surface temperature (°C), 0.5 m below the surface. T bot = Bottom temperature (°C), 0.5 m above the sediment. ΔT = Stratification strength (°C) (= T surf−T bot). P atm = Atmospheric pressure (Pa). Depth = Water column depth (m). ΔP tot/Δt = Gradient of total pressure (atmospheric + hydrostatic) (Pa h−1). Wind = Wind speed (m s−1). O 2,bot = Oxygen concentration 0.5 m above the sediment (mg L−1). Chl‐a = Chlorophyll‐a (μg L−1). NEP = Net ecosystem production (gO2 m−3 d−1). GPP = Gross primary production (gO2 m−3 d−1). Inflow = Inflow discharge (m3 s−1). B 0,net = Net surface buoyancy flux (W kg−1). SWR = shortwave radiation (W m−2).
Figure S9: Reservoir‐wide total emissions. (a, b) Reservoir‐wide cumulative (a) CH4 and (b) CO2 emissions, expressed in moles and assuming that the EC footprint is representative of emissions from the entire reservoir, and (c) reservoir surface area during the wet and dry years.
Figure S10: Random forest predictions for the original and decomposed CH4 flux signals during the dry year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S11: Random Forest predictions for the original and decomposed CH4 flux signals during the wet year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S12: Random Forest predictions for the original and decomposed CO2 flux signals during the dry year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S13: Random forest predictions for the original and decomposed CO2 flux signals during the wet year. Panels show 1:1 relationships (a, c, e, g) and corresponding time series (b, d, f, h) for the observed signals and the Random Forest (RF)–predicted signals. Black lines in panels (a, c, e, g) indicate the 1:1 relationship. For each predicted signal, the number of predictors (N), the selected predictors, and the coefficient of determination (R 2) are shown. The five most important predictors were initially selected based on their permutation‐based out‐of‐bag (OOB) importance, and further predictor reduction was applied when R 2 exceeded 0.95 using a smaller subset of predictors.
Figure S14: Bottom water temperature vs. methane ebullitive and diffusive fluxes. The black dashed line shows the best 2‐year Arrhenius fit [Equation (1)], with the corresponding activation energy (E A ) indicated. The gray shaded area around each Arrhenius fit represents the 95% confidence interval.
Figure S15: Random forest predictor importance for the decomposed diurnal diffusive (a) and ebullitive (b) CH4 fluxes. Diffusive and ebullitive components were extracted from the total eddy covariance CH4 fluxes using the wavelet analysis approach proposed by Iwata et al. (2018). The decomposed temporal flux components were obtained using the maximal overlap discrete wavelet transform (MODWT) and multiresolution analysis. Each Random Forest analysis was repeated 50 times using different random seeds to assess model stochasticity. Bars represent mean predictor importance across runs, and black horizontal lines indicate ±1 standard deviation.
Figure S16: Daytime versus nighttime fluxes. Daily‐averaged gap‐filled CH4 (a, b) and CO2 (c, d) fluxes during daytime (PAR ≥ 10 μmol photons m−2 s−1) and nighttime (PAR < 10 μmol photons m−2 s−1) conditions for the dry year (a, c) and the wet year (b, d).
Figure S17: Hydrostatic pressure versus methane fluxes. Relationship between daily‐averaged hydrostatic pressure at the lake sediment surface and methane fluxes. Solid gray lines show the seasonally decomposed relationship between the two variables, and arrows indicate the direction of progression over the annual cycle. Linear regressions are shown for periods during which CH4 fluxes increase linearly with decreasing hydrostatic pressure (i.e., water depth). The gray shaded area around each regression represents the 95% confidence interval.
Table S1: Coefficient of determination (R 2) for random forest predictions as the number of variables (within brackets) increases from 1 to 5, or until a combination reaching R 2 > 0.95 is achieved. Variables 1–5 were selected based on their predicted importance (the most important first).
Data Availability Statement
The data supporting the findings of this study are openly available in https://doi.org/10.5281/zenodo.20800380.
