Figure 1(c) shows the climatology of vegetation productivity of mangroves. Vegetation productivity in this region follows a bell-shaped curve, with maximum productivity during the summer monsoon (June to September, known as the Indian Summer Monsoon or ISM), when the Indian subcontinent receives 80% of its annual rainfall52–55. The study area also receives some precipitation due to local thunderstorms called ‘Norwester events’56 in pre-monsoon. This is reflected as a slight jump in vegetation productivity in April-May in Fig. 1(c). These events bring intense rain and strong winds and can last a few hours. The period from December to March generally has low vegetation productivity as there is limited freshwater due to minimal precipitation. Supplementary figure S1(a-e) shows the anomalies (diurnal cycle removed) of eco-hydro-meteorological variables from January 2013 to December 2015 averaged to a weekly time scale. The figure shows all variables plotted with NEE as the second axis to visualize their variations with carbon fluxes.
Winters (figure S1) are characterised by lower TA and P, leading to lower LE and high VPD. During this period, WS is also low. During pre-monsoon, LE rises due to increased evapotranspiration under high TA. P during pre-monsoon is scarce and mainly occurs in the form of thunderstorms. During pre-monsoon, WS is high, and VPD starts reducing as air holds more moisture because of more land evaporation and moisture brought by hot air from the Bay of Bengal. During monsoon, LE drops as TA is reduced due to high P and continuous cloud cover. Winds during monsoon are strong, and VPD is very low because of the high moisture content of the air and continuous rainfall. The high LE in 2014 (figure S1(a)) is because of the strong El-Nino of 2014-15, which reduced P and high TA during ISM57,58. The presence of El Nino years in our three-year study period will change the location and scale parameters of the distributions; however, due to the limited availability of data, we decided to choose this study period.
Stability against changing hydro-meteorological variables
Figure 2(a-d) shows standardized deviations of daily averaged flux tower data from respective monthly means plotted as three-day moving averages for NEE, P, WS, and TA, respectively. For example, the mean value of January month (90 values) is subtracted from all values of January in the daily time series, and thus generated anomalies for January days are divided by the standard deviation of January month (90 values). We show three-day moving averages instead of daily anomalies to smoothen the signal for better visibility. Dotted lines in the plot represent thresholds of data being one standard deviation away from the mean. While hydro-meteorological variables show significant variation throughout the observation period, generating values greater than one standard deviation, NEE remains mostly stable. For measuring this stability, we defined the Mangrove stress period as the period for which the Mangroves have NEE continuously greater than one standard deviation from the mean value. We fitted the beta distribution to the duration of this stress period (using daily data, histogram shown in Fig. 2f, and best-fit distribution Fig. 2e) and found that the average duration lies between 1–2 days with 95% confidence (statistically significant using the sample of size 91). The low mean value of the distribution shows a rapid recovery of NEE from perturbations with an average recovery period of between 1 and 2 days.
Stability against deteriorating nutrient supply
Figure 3 shows boxplots of NEE (Fig. 3(a)) from the flux tower during the study period (2013–2015), along with the observed monthly in-situ measurements of nutrients. Figure 3(b-d) shows Nitrate (Ni) to Phosphate (Ph) ratio (N/P), silicate (Si), and Ammonium (Am), respectively. Coastal areas have an ideal ratio of Si to Ni to Ph as 106:16:1, called the Redfield ratio49,59, which is essential for ensuring optimal vegetation productivity. We found that the N/P ratio has constantly increased from being close to the ideal value of around 16:1 in 2013 to an average value of around 75:1 in 2015. The increase is roughly five-fold, with a few months showing around 10-fold increases from 2013 to 2015 (Fig. 3b). During this period, the amount of Ni increased, and Ph decreased continuously from 2013 to 2015. A decrease in P indicates the phosphorous limitation on vegetation productivity, which has previously been reported in the literature40. Si and Am have also shown an increase of around 3 times (from 20 µM to around 60 µM average) and 7 times (from 1 µM to around 7 µM average), respectively. We found that while the nutrient composition has deteriorated significantly during the study period (Fig. 3(b-d)), NEE has remained stable (Fig. 3(a)). No apparent change in mangrove NEE is an interesting observation as it indicates that mangroves have so far been able to resist anthropogenic stressors which are known to reduce vegetation productivity by impacting plant physiology across short-term and long-term levels. The stability of mangroves becomes even more surprising because the changes in nutrients were also accompanied by a prolonged El Nino event, which is known to reduce the vegetation productivity and cause droughts across South Asia57,60.
Response mechanism through varying process connectivity
The aforementioned findings raise the following query: how do mangroves stabilize their productivity under varying natural and anthropogenic stressors? To answer this, we examine the changes in process connectivity of mangrove NEE with hydro-meteorological variables through weekly and monthly causal networks generated using PCMCI (see methods). Since PCMCI can delineate non-linear causal associations, the temporal evolution of causal networks thus generated should reveal the evolving dynamics of mangroves under anthropogenic stressors. Figure 4 (a) shows the mean incoming link strength to NEE in the weekly networks. Vertical lines indicate the beginning and end monsoons (uniformly taken as 1st June to 30th September) of three years. We find that the incoming link strength from other variables to NEE has increased from 2013 to 2015 (the statistically significant increasing trend at 95% confidence using 158 data points). The increase in the connection strength is also accompanied by an increase in lags (Fig. 4(b), statistically significant increasing trend at 90% confidence using 158 data points) which indicates that physiological processes of mangroves are becoming more memory-driven under anthropogenic stressors. As an example, we show the networks of December month generated using 3-hour data (248 values) for two years, 2013 and 2015 (Fig. 4 (c) and (d), respectively). For December, incoming link strength to NEE from TA and P became stronger, and one new link from VPD was established, while links from LE and WS were lost. More figures for each month of each year are presented in supplementary figure S2. The increase in the average incoming link strength from 2013 to 2015 is clearly visible for most of the months; however, the increases cannot be narrowed down to a single source variable. The connections are dynamic and vary from month to month. While some links disappeared from 2013 to 2015, new links appeared with stronger connection strengths and larger lags. Hence, the mangroves are able to resist and adapt to increasing anthropogenic stressors by reorienting the strength and memory of their dynamic process connectivity with hydro-meteorological variables.
Process connectivity during weather extremes
To delineate the process changes in mangroves during weather extremes, we analyzed weekly causal networks around the cyclone Viyaru, which crossed the study area from 10th – 14th May, 201361, and a norwester event in April 2013. We present our results in Figures S3 and S4, respectively. For cyclone (Figure S3), we showed five weekly networks starting from 29th April 2013, which include the cyclone at the end of the second week. From the number of incoming links to NEE in the networks, we clearly see that the incoming links to NEE drop immediately after the cyclone (figure S3(c)) and quickly recover in the next week (figure S3(d)) and a further increase in the following week (figure S3(e)). We found a similar case for the norwester event, as shown in figure S4. We show the number of incoming and outgoing links with node NEE for five weeks with a ‘norwester’ event lying at the third week (figure S4(a)) to include two preceding and two succeeding weeks in the analysis. We found that incoming links to NEE drop during the week of the thunderstorm but quickly recover in the next week. Outgoing links from NEE do not show the same signature as they represent the feedback from vegetation to the atmosphere, which might depend on other factors and is still poorly understood at short time scales. Overall, we found that mangrove NEE is stable to short spells of extreme weather events like thunderstorms. It shows that dynamic processes involving NEE in mangroves are briefly disturbed during weather extremes but are restored within one to two weeks.