Impacts of MSP on seedling survival in interaction with summer drought
Our study showed that MSP enhances first-year seedling survival of species used to a large extent in European planted forests (sessile oak, Douglas fir, several pine species). It also showed that the increase in seedling survival following MSP is more pronounced when the summer is dry. These observations support our first and second hypotheses and agree with a large body of literature (Löf et al., 2012).
In years with rainy summers, early seedling survival in plots without MSP was high. MSP brought a slight but significant improvement and was similar for moderate and severe MSP. Seedling survival rates in plots without MSP decreased sharply as the amount of summer rain decreased, as previously shown by Zwolinski et al. (1994). In dry summers, seedlings planted after severe MSP had higher survival rates than those planted without MSP or with moderate MSP. All together, these observations partly invalidate our third hypothesis since severe MSP was more beneficial to seedling growth than moderate MSP only in dry summers, and they support our fourth hypothesis since the benefit from severe MSP, compared to moderate MSP was higher in years with summers drier than normal.
The novelty of our study was to analyze the interaction between MSP severity and drought intensity. In rainy years, moderate MSP resulted in the same seedling survival rates as severe MSP, whereas in dry years, moderate MSP led to the same rates as the control without MSP. The interaction we observed between MSP severity and drought intensity suggests that moderate and severe MSP not only differ in the severity of their impact on the seedling environment but also in the type of impact they induce. MSP may improve seedling survival through the reduction of competition with neighboring vegetation, insect or rodent damage, soil compaction or soil water logging, and the various MSP methods differ in their impacts on these constraints (Löf et al., 2012). The relative importance of the constraints for seedling survival depend on the climatic conditions, soil characteristics and biotic interactions prevailing in the plantation site, and the impact of a MSP method on seedling survival in a specific set of environmental conditions may be indicative of which constraints are primarily affected by the MSP method.
In our study, moderate MSP was effective at reducing constraints that limited seedling survival in years with rainy summers (competition for nutrients and light from neighboring plants, biotic damage mediated by neighboring plants, allelopathy caused by neighboring plants, soil constraints), but could not significantly reduce constraints that limited seedling survival in years with dry summers (the same constraints as above, plus competition for water from neighboring plants). Constraints not related to drought were probably limited, as suggested by the magnitude of the gain in survival brought by MSP in rainy years (+ 7%). These constraints probably also occurred in dry years as well but could not be detected because they were small compared to the high variability in survival observed in these years. This observation may explain the absence of statistically significant effects of moderate MSP in dry years, compared to the unprepared control. A larger number of observations would be required to test their effect.
Severe MSP was as effective as moderate MSP to reduce constraints that limited seedling survival in rainy years. In addition, severe MSP significantly increased survival rates in dry years, showing that severe MSP efficiently alleviated drought constraints (i.e., mainly competition for water from neighboring plants).
Methodological approach
Our study followed an experimental approach to analyze the combined effects of MSP and drought intensity on seedling survival. In the place of an experimental network, previous studies (Boutte et al., 2023; Tallieu et al., 2022) used large-scale inventories of forest plantations managed by forest operators to examine the same effects as our study on a similar geographical area (entire France), a similar time period (2017–2021) and a list of tree species and MSP methods that included the species and MSP methods tested in our study. However, although they used a large dataset (more than 10,000 plantation sites), they did not observe any significant effects of MSP on seedling survival, once potential confounding factors (species, other silvicultural operations, geographical area) were taken into account in the models. In this work, the lack of response was ascribed to (1) correlations between the predictive factors throughout the dataset since, in practical forestry, MSP methods are selected according to the silvicultural context; and (2) strong variability in seedling survival observed within each level of the predictive factors. The experimental approach used in our study made it possible to overcome these limitations: (1) we balanced the tree species and the MSP methods along the geographical gradient explored by the experimental network, which successfully led to weak correlations among our predictive factors; and (2) the establishment of an unprepared control in each experimental site made it possible to pair MSP and control treatments, which absorbed a strong proportion of the observed intra-treatment variability in seedling survival. The approach brought a high statistic power to the analysis and allowed us to observed significant effects of MSP in interaction with drought intensity, using a relatively small number of experimental sites.
As expected from previous work in the field of disturbance ecology (Peterson and Anderson, 2009; Royo et al., 2016), the severity of impact was successfully used to compare disturbances (in our study: MSP methods) with distinctive features. We analyzed nine MSP methods that differed in the type of tool and prime mover used, and in the type and the spatial extent of the preparation performed. The severity index made it possible to synthesize their features into a single descriptor that could successfully predict the effects of MSP on planting success.
We selected the cut-off value to distinguish between moderate and severe MSP methods in order to obtain a sufficient number of observations in each severity class, for the sake of the statistical analysis, without any ecological or technical consideration. Although our study led to significant results, further work could be performed to test different cut-off values and to examine their ecological or technical significance in order to improve our global understanding of the processes that underlie seedling response to MSP.
Although MSP is known to affect several environmental factors in the immediate seedling vicinity (e.g., water, light and nutrient availability, soil features, susceptibility to biotic damage), we used a single indicator to estimate MSP severity, which was the vegetation cover one year after MSP. In the field of disturbance ecology, combining several indicators that reflect the different ecosystem processes affected by disturbances provides a consistent framework for analyzing, comparing and modeling the effects of disturbances on ecosystems (Roberts, 2007). Although different indicators are usually positively correlated along gradients of increasing severity, substantial variations may occur among indicators, and combining several indicators may help elucidate the factors and mechanisms that underlie ecosystem responses to disturbance (Roberts, 2004).
In our study, we used a single indicator of MSP severity: vegetation cover in the immediate seedling vicinity one year after MSP. It performed well because (1) following MSP, vegetation cover positively correlates with other indicators of MSP intensity such as the extent of soil disturbance (Chaves Cardoso et al., 2020); (2) vegetation control is one of the primary effects of MSP (Löf et al., 2012); (3) in our study, vegetation cover could be estimated with good accuracy; and (4) in our study, we explored a large gradient of vegetation cover values (ranging from 3 to 100%). In a preliminary analysis, we also used the amount of soil disturbed by the MSP method (estimated through the maximum soil depth reached by the MSP tool and through the surface area of soil disturbed), as suggested by the previous study of Sikstrom et al. (2020). However, the soil indicators ranged along a small gradient of soil disturbance values (due to the selection of MSP methods in our study) and were difficult to estimate accurately (due to the measurement methodology in our study). Eventually, the soil disturbance indicators did not further discriminate between MSP methods, and we eliminated them from further analysis and instead used a single indicator based on vegetation cover. Selecting a different set of MSP methods that showed more pronounced differences in soil disturbance could possibly have led to different conclusions on the efficiency of the soil indicators used to estimate MSP severity.
Recommendations for management
We observed a strong decrease in seedling survival when summer drought increased, which is in full agreement with the long-term monitoring of plantation success in France (Boutte et al., 2023; Tallieu et al., 2022). In our study, we were not able to observe species-specific responses due to the small number of experimental sites available but, in their long-term monitoring, Tallieu et al. (2022) showed that some species (including Quercus petraea and Pseudotsuga menziesii.) are very responsive to summer drought, while others (including Pinus sylvestris, Pinus nigra and Pinus pinaster) show constant survival rates irrespective of summer drought. These observations provide robustness to our results since our results apply to tree species that show a wide range of responses to summer drought. Similarly, due to the limited number of experimental sites, we were not able to observe any impacts of vegetation or soil type, although these factors are known to have primary effects on seedling response to MSP (South et al., 2023), and forest managers should use their own expert knowledge to translate the results of our study to specific site conditions.
In many European countries, 1-year-survival is used as a criterion to evaluate planting success, with a threshold value set between 80 and 90% (Mataruga et al., 2023). In France, in most reforestation contracts, a threshold value of 80% seedling survival is used, below which the company that was responsible for the plantation is committed to paying for all refilling operations. We will use this threshold value as a basis to discuss practical recommendations based on our results.
In our study, the average survival rate in plots without MSP was 50% in dry years, which is well below the threshold value of 80%. In rainy years, seedling survival was above the threshold of 80% in most plots without MSP (13 out of 17 plots). All together, these observations reaffirm that the main issue for forest managers in France is to ensure plantation success during dry years for species that are most susceptible to summer drought, in order to avoid the extra costs due to plantation refill.
In rainy years, a positive and statistically significant effect of MSP on seedling survival was observed, irrespective of MSP severity. However the magnitude of the effect was small (+ 7%) and its ecological or practical significance (sensus Dixon and Pechmann (2005) and Wei et al. (2015)) is questionable. Defining a threshold value above which the effect of MSP may be considered as practically significant requires expert knowledge and data on long-term plantation dynamics and economic value that were not available for our study. Previous studies performed on Pinus taeda L. (South et al., 2023) suggest that the benefit of MSP during the rainy years that we observed in our study was too small to substantially increase stand value at harvest and, therefore, could be considered as negligible.
In dry years, severe MSP made it possible to significantly increase seedling survival compared to plots without MSP, and the magnitude of the effect (+ 30%) is large and, most probably, significant for the forest practice. Only three plots out of 11 that received severe MSP showed a survival rate below the threshold value of 80%. In practical forestry, these three plots would be refilled, and the eight remaining plots would avoid the expense of refilling operations. In comparison, eight out of nine plots in the control and seven out of eight plots in the moderate MSP treatment were below the threshold value and would need to be refilled. Applying a severe MSP method reduces the number of plantations to be refilled after a dry summer by 60%, compared to applying a moderate or no MSP. However, we can observe that, even if severe MSP significantly improves seedling survival, it does not guarantee obtaining a survival rate above the 80% threshold in dry years.
MSP has a financial cost that must be balanced against its beneficial effect on seedling survival. In our study, severe MSP methods used tools mounted on medium sized (7 to 8 tons) or large (20 to 22 tons) excavators, whereas moderate MSP methods (except for treatment #2) used tools pulled by tractors. Using an excavator instead of a tractor approximately doubles the working time (Puyal et al., 2022) and the financial cost (Suadicani, 2003) of the preparation. In rainy years, there is no benefit to using excavators that perform severe MSP instead of tractors that perform moderate MSP because they are more costly and do not improve early seedling survival. On the contrary, in dry years the benefit of using excavators may be high because they increase seedling survival, although they are more costly. In each reforestation project, the MSP method is chosen in the autumn or the winter prior to planting, whereas the potential benefit on seedling survival can only be seen several months later during the summer that follows planting. Determining, prior to planting, whether a MSP is required and choosing the most appropriate MSP method for each plantation project would require a risk analysis that takes into account (1) the probability of summer drought occurrence; (2) the probability of seedling survival according to drought intensity and the MSP method; (3) the cost of the MSP methods; and (4) their long-term economic return as a function of initial seedling survival. A research effort to gather such data is presently required for most plantation systems in order to develop decision support tools that make it possible to choose the best MSP method in each plantation site. Meanwhile, before such practical tools are available, forest managers may secure their plantation projects in dry and in rainy years by applying a severe MSP or by opting for an insurance contract against plantation failure (Bréteau-Amores et al., 2023).
Finally, there is presently a growing emphasis on the maintenance of soil fertility and biodiversity, prompting forest managers to select silvicultural methods that preserve both soil and biodiversity (Duncker et al., 2012), and we also need to consider environmental impacts of MSP, when recommending appropriate MSP methods. In our study, MSP severity was assessed in the immediate seedling vicinity (1-m2-subplots surrounding the seedlings), which corresponds to the rooting zone of the newly planted seedlings and the potentially competing neighboring plants. We found that MSP severity was related to seedling survival. We must stress that our results do not suggest that implementing a MSP method beyond the area immediately around the seedlings would further improve seedling survival. Therefore, implementing MSP on a larger surface area (along the planting line or throughout the entire stand area) should be decided cautiously, given its demonstrated strong and long-lasting environmental impact, particularly on soil structure and fertility (Collet et al., 2020; Sutinen et al., 2019). In this respect, intermittent or spotted MSP methods (Sikström et al., 2020) that implement a severe preparation around the planted seedlings while leaving the rest of the surface area in the stand undisturbed, appear to be a good option for forester managers looking to ensure high seedling survival while simultaneously preserving the environment.