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Two potential equilibrium states in long-term soil respiration activity of dry grasslands are maintained by local topographic features

Spatial patterns of stability proxies and background variables

Figure 2 a, b show the spatial distribution of our two proxy variables, the average rank of Rs per position (rankRs) and of the range of the ranks per position (rangeRs) in kriged maps. The middle to southern areas were found to have the largest, whilst the north-eastern areas the smallest rankRs values, whereas a slightly different pattern was characteristic for rangeRs with some additional north-western large values. Similarly, larger average soil organic carbon content (meanSOC) and average soil water content (meanSWC) (Fig. 2 c, d) were detected at the western-middle-southern regions and smaller at the north-eastern part of the study site.

Figure 2

Kriged patterns of stability proxies, rankRs (a) and rangeRs (b), as well as of background factors, meanSOC (%, c) and meanSWC (%, d).

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Correlations between stability proxies and background variables along DEMs: entire dataset

We investigated the potential direct effects of the different terrain attributes (local mean elevation (mALT), standard deviation of elevation (SD), topographic position index (TPI), slope (Sl), Easterness and Northness (East, North)) on the spatial distributions of our proxy variables by using the terrain attributes originating from differently smoothed DEM rasters. DEM1 was the original, 0.2 m resolution model, while DEMs 2–6 were progressively smoothed by a factor of two resulting in different resolution DEM rasters (DEM2: 0.4 m, DEM3: 0.8 m, DEM4: 1.6 m, DEM5: 3.2 m, DEM6: 6.4 m, respectively), and finally DEM7 met the resolution of the field measuring campaigns (10 m). The terrain attributes were filtered out from the rasters for the 78 measuring positions of the sampling grid.

On the basis of the correlation analysis we found an important difference in terrain attribute features between DEM 5 and 6, especially in SD, Sl, North and East. All subsequent results are then based on DEMs 1–5, which were found to be more similar to each other and to the original DEM1. The maps of terrain attributes with the box blur kernel from DEM1-5 can be found in the Supplementary Information (SI) together with the descriptions and calculations. As we couldn’t find any of the blur kernels superior to the other when considering correlations, the results hereafter are only presented for the box blur kernel calculations for simplicity.

When we considered the entire dataset (named hereafter: “A” dataset), we could only find significant correlation between rangeRs and TPI at less smoothed DEMs but the correlation was very weak (black symbols and line in Fig. 3).

Figure 3

Direct correlation between TPI and stability proxy, rangeRs at less smoothed DEMs, DEM1-2 for datasets A (black symbols and line) and S (blue symbols and line, see the information later on). The correlations were significant at p = 0.0076 and p  = 0.0172 levels, although they were weak, r2 = 0.09, r2 = 0.42 for A and S (see the information later on), respectively.

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Any other correlation between the proxies and the terrain attributes could only be deduced indirectly from the positive correlations between rankRs and meanSOC, meanSWC (cf. Table 1b). These correlations were scale-independent, i.e., we detected them at every DEMs. In general, the larger the soil carbon content and soil moisture at a position (cf. Figure 2c,d, showing quite similar patterns to the proxy patterns in the figure upper row), the larger the Rs activity detected and the opposite was true for lower carbon content and soil moisture positions.

Table 1 (a) Statistically significant (p < 0.05) linear correlation between terrain attributes, ALT, mALT, TPI, SD, Sl, North, East and background factors, meanSOC, meanSWC for A dataset. (b) Statistically significant (p < 0.05) linear correlation between terrain attributes, background factors and stability proxies, rankRs, rangeRs in A dataset and in the subgroups (see codes in the text). Regular letters mean scale-independent correlations (valid for DEM1-5), italic underlined letters mean correlations valid for less smooth DEMs (DEM1-2 or 1–3), “pos” and “neg” indicate the sign of the correlation.

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As we investigated the background of these correlations more thoroughly in dataset A in terrain attributes (cf. the maps in SI, Table 1a), we found that meanSOC showed negative correlation with ALT, mALT, SD and Sl, while positive with North and East at DEMs 1–5. Similarly, meanSWC correlated negatively to SD, Sl, except for DEM5. Several terrain attributes were then responsible for the meanSOC patterns, i.e., higher absolute elevation and neighbouring surface heterogeneity, as well as steeper slope positions facing more South-West could be characterized with lower meanSOC. The opposite features were characteristic for higher meanSOC level positions on lower elevations with lesser neighbouring heterogeneity and gentle slopes facing mostly North and East. Similarly, meanSWC was higher at smaller surface heterogeneity with more gentle slopes, while lower at more heterogeneous surfaces with steeper slopes. rankRs followed these patterns with higher Rs activity in the middle-southern part of the study area, while, in contrast, lower Rs activities were characteristic at lower meanSOC and meanSWC at the north, north-east facing locations in the study site, mainly on local ridges as found on the basis of the direct TPI correlations.

Correlations between stability proxies and background variables along DEMs: subgroups

We also checked the correlations within different data subgroups corresponding to specific rankRs or meanSOC categories because we hypothesized that these kinds of groupings could enable us to grasp some important characteristics of the stability.

Subgroups:

  • Subgroups were created on the basis of rankRs ± SD: S (Smaller than mean-sd), M (Middle between mean ± SD), L (Larger than mean + SD).

  • C1, C2, C3, C4, C5 (from the smallest to the largest meanSOC quintiles).

Direct correlation between terrain attributes and proxies showed considerable variation depending on the subgroups and variables (Table 1b collects scale-independent correlations, valid at almost each of the DEMs between 1 and 5, or scale-dependent ones, valid only in several of the less smoothed DEMs 1, 2, 3).

It seems that the meanSOC pattern related negatively to ALT, mALT detected in dataset A could have acted as a driver for the negative rankRs and ALT, mALT correlations in the groups M and C1. It was in subgroup C1 that TPI, SD and Sl acted negatively on rankRs as well, most probably more directly through the patterns generated by terrain attributes in meanSWC. Further negative correlations were found between rangeRs and TPI in S data (see also blue symbols and line in Fig. 2), like in dataset A (cf. Fig. 3 black symbols and line), as well as between rangeRs and North in C4. Accordingly in the long run, local valleys but mostly constant slope positions (with TPI close to zero, cf. blue symbols in Fig. 3) with lower neighbouring surface heterogeneity and gentle slopes with more elevated meanSOC and meanSWC could be characterized with larger Rs activity with higher variability (through the negative TPIrangeRs correlation) in these subgroups per se, similarly to dataset A. The opposite was likely to be the case for local ridges.

The subgroups mentioned here were restricted groups of measuring positions, where carbon content in the soil was the lowest of all (C1) or, as in subgroup S, coincided with low meanSOC levels (Fig. 4), and low meanSWC levels as well.

Figure 4

Density plots of meanSOC and meanSWC while grouping the measuring positions according to their rankRs category (S-M-L groups) or meanSOC content (C1–C5 groups).

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Furthermore, measuring positions, grouped by either on the basis of rankRs or on the basis of meanSOC, occupied more or less well delimited spatial areas within the sampling grid (Fig. 5, positions coloured according to C1–C5, where e.g., C5 category, indicated with asterisk occupied the lowest altitudinal positions, C4 was found mostly around C5, while C1 category could be found along the edges of the study area on the crests), which would also be characteristically different in respect of the terrain attributes, especially in SD, Sl, as found by the correlations (cf. Table 1).

Figure 5

Digital elevation model of the study plot in Bugac, Hungary (coordinates refer to the Uniform National Projection System (m)) with the sampling positions in the 80 × 60 m grid coloured according to their meanSOC from C1 to C5. Green square represents the position of the eddy covariance station. Marginal plots in grey show the mean surface elevation by x and y coordinates. Notice that the largest altitudinal difference was no more than ~ 1.5 m within the study plot.

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Finally, rangeRs was fitted to rankRs using the following equation:

$$rangeR_{s} = a times frac{1}{{sigma sqrt {2pi } }}e^{{ – left( {rankR_{s} – mu } right)^{2} /2sigma^{2} }} ,$$

(1)

where µ is the mean rankRs (42.61), σ is the standard deviation of rankRs (25.38), and a (4,291.73) is a model parameter. The correlation, approaching a bell-shaped curve (Fig. 6, both the curve and the model parameters are statistically highly significant, p < 0.0001), visualized together with the subgroups showed that both low and high rankRs could be associated with small rangeRs with larger stability and a typically resistant response, while middle values of rankRs corresponded to larger rangeRs with a more flexible, resilient response of Rs activity. Furthermore, it was also showed that rankRs categories more or less fitted to C1–C5 meanSOC categories (cf. square symbols of C1 in S-M-group regions, C2 in M, while asterisks mostly in the upper half of the rankRs range), giving strong evidence of SOC as a controlling factor in Rs stability. The smallest and the largest rankRs values could correspond to the largest potential stability (in terms of resistance) in the activities, rankRs being either low in general (cf. Fig. 2a north and north-east regions) due to low meanSOC and meanSWC (Fig. 2c,d) or high (cf. Figure 2a more the middle and southern regions within the study plot) in the opposite cases. On the other hand, medium rankRs with larger rangeRs overlapped spatially with C2-C3 and M groups with medium meanSOC levels, and these positions showed a more resilient response.

Figure 6

Bell-shaped curve correlation between rankRs and rangeRs visualized together with the two series of subgroups, S-M-L created on the basis of rankRs categories, C1–C5 created on the basis of meanSOC categories. Equation and parameters of the curve are in the main text.

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