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All the environmental conditions necessary for natural forest regeneration constitute a favorable microsite for seed germination and seedling growth. Among these microsites, terricolous bryophytes represent an important support, present in almost all ecosystems. In addition, bryophytes have an important role in the recruitment of certain vascular plants in temperate forests. Terricolous bryophytes, through the expression of certain morphological traits such as density or mat thickness, can modify regeneration microsites by modulating soil temperatures; moisture and access to nutrients (filter theory). However, the interactions between tree forest regeneration and bryophytes remain poorly understood. In order to understand the interactions between bryophytes and regeneration, we studied the general effects of vegetation strata and bryophytes on natural forest regeneration in a causal system (Structural Equation Modelling) on a national network of observation plots. In a second step, we conducted a regeneration monitoring in mixed forest on a regional network of experimental plots to identify the effects of bryophytes on the growth and survival of sessile oak (Quercus petraea) and Scots pine (Pinus sylvestris) seedlings. We also tried to evaluate the effects of two functionally different bryophyte species on the abiotic conditions of the microsite by monitoring temperature, humidity and light in situ. Finally, we conducted a semi-controlled greenhouse experiment to monitor germination and growth of oak and pine on bryophyte mats as well as on bare soil as a function of two atmospheric and two edaphic humidities. We show and discuss that, depending on their functional traits; different bryophyte species do not have the same effect on seedling establishment, growth, or survival rates. Bryophytes may have a physical barrier effect but also play a role on moisture in regeneration microsites. Terricolous bryophytes would filter out different tree species depending on their seed size., L’ensemble des conditions environnementales nécessaires à la régénération forestière naturelle constitue un microsite favorable pour la germination des graines et la croissance des semis. Parmi ces microsites, les bryophytes terricoles représentent un support important, présent dans la quasi-totalité des écosystèmes. De plus, les bryophytes ont un rôle important sur le recrutement de certaines plantes vasculaires dans les forêts tempérées. Les bryophytes terricoles, par l’expression de certains traits morphologiques comme la densité ou l’épaisseur du tapis, peuvent modifier les microsites de régénération en modulant les températures, l'humidité et l’accès aux nutriments du sol (théorie des filtres). Cependant, les interactions entre la régénération forestière arborée et les bryophytes restent méconnues. Afin de mieux comprendre les interactions entre les bryophytes et la régénération, nous avons étudié les effets généraux des strates de végétation et des bryophytes sur la régénération forestière naturelle dans un système causal (Modèle d’Équations Structurelles) sur un réseau national de placettes d’observation. Dans un deuxième temps, nous avons réalisé un suivi de régénération en forêt mélangée sur un réseau régional de placettes expérimentales pour identifier les effets des bryophytes sur la croissance et la survie des semis de chêne sessile (Quercus petraea) et pin sylvestre (Pinus sylvestris). Nous avons également essayé d’évaluer les effets de deux espèces de bryophytes fonctionnellement différentes sur les conditions abiotiques du microsite en réalisant des suivis de température, d’humidité et de lumière in situ. Enfin, nous avons réalisé une expérimentation en conditions semi-contrôlées en serre pour suivre la germination et la croissance du chêne et du pin sur des tapis de bryophytes ainsi que sur le sol nu, en fonction de deux humidités atmosphériques et de deux humidités édaphiques. Nous montrons et discutons qu’en fonction de leurs traits fonctionnels, les différentes espèces de bryophytes n’ont pas le même effet sur l’installation, la croissance ou le taux de survie des semis. Les bryophytes peuvent avoir un effet de barrière physique mais jouent également un rôle sur l’humidité dans les microsites de régénération. Les bryophytes terricoles filtreraient différentes espèces arborées notamment selon la taille de leur graine.

Accord par mail pour la mise en ligne du pdf éditeur; National audience; Les mélanges d’essences et la baisse de la densité des arbres dans les peuplements sont des pratiques envisagées comme solution face au changement global. Elles peuvent aider à préserver les forêts et les services qu’elles rendent comme la production de bois, le stockage de carbone et la préservation de la biodiversité. Le dispositif OPTMix, unique en France, étudie les effets du climat, de la gestion sylvicole (mélange et densité) et de la présence des grands herbivores, et leurs interactions sur le fonctionnement des forêts de plaine. Réparti sur quarante hectares, il comporte trente-trois placettes de chêne sessile (Quercus petraea) et de pin sylvestre (Pinus sylvestris) en peuplements monospécifiques ou en mélange, fortement instrumentées pour mesurer le microclimat, la croissance des arbres, la dynamique du sous-bois dont la régénération du peuplement, la gestion des ressources (eau, lumière, nutriments), et la biodiversité (flore vasculaire, bryophytes, lichens, carabes, microfaune du sol). Les premiers résultats montrent que l’intérêt du mélange n’est pas systématique pour les différents services étudiés, et aussi que l’identité des essences détermine à la fois l’intensité et le sens des effets observés. Les dernières années de fortes sécheresses montrent également que l’intérêt des mélanges reste limité pour faire face aux changements climatiques annoncés. De plus, pour la biodiversité, il est recommandé de conserver une mosaïque de type de peuplements à l’échelle du massif, avec des purs de différentes essences et des mélanges plutôt que de généraliser les peuplements mélangés.

In most mixed-species forestry systems, regeneration is a critical step for individual tree survival and species coexistence. Species coexistence is driven by the fine-scale spatial patterns and competitive abilities of species. In naturally regenerated mixed oak (Quercus petraea and Quercus robur: QP and QR, respectively) stands, species coexistence was assessed by analysing the spatial distribution of both species and their growth responses to intra- and interspecific competition. Four sites located in North Eastern France with an established (1.4–2.8-m-high) mixed QP and QR regeneration were selected for the study. In a first step, individual seedling location was recorded along line transects to analyse the spatial distribution of the two species. Univariate pair correlation functions were used to test for spatial aggregation for each species and bivariate pair correlation functions were used to test for spatial associations between the two species. In a second step, seedlings were measured in small plots to analyse their response to local competition. Mixed linear models were used to estimate the combined effects of density, mixture, seedling size and seedling status on seedling growth. QP and QR were both highly aggregated in monospecific patches, and the two species were strongly segregated at a decametric scale. Transition zones between monospecific patches were reduced to a few meters. For both species, mixture had no effect on seedling growth. In all sites, the two species showed very similar diameter and height growth rates in response to density, mixture, seedling size and seedling status, and similar intra- and interspecific competitive abilities. Highly patchy patterns of QP and QR regeneration strongly reduce interactions between the two species and, in transition zones between monospecific patches where the two species are mixed, no species is expected to outcompete its companion species. For each species, long-term persistence at the stand scale is determined by intraspecific competition rather than by interspecific competition. At this developmental stage, the QP-QR mixture appears to be very stable and no specific silvicultural treatment is required to maintain the mixture.

L'étude du fonctionnement des peuplements mélangés n'est pas près de s'épuiser. En ce qui concerne l'effet du mélange sur la productivité, les travaux ont surtout porté sur la croissance radiale ou en surface terrière. Mais la productivité dépend aussi de l'influence du mélange sur la croissance en hauteur, plus difficile à appréhender : les deux effets sont-ils de même ampleur ? Vont-ils dans le même sens ? Quels sont les mécanismes ? Méthode d'étude et premières réponses.

International audience; Coppice forestry is a conventional silvicultural practice that takes advantage of a tree's capacity to respond to disturbances by sprouting. Sprouting capacity is determined by many intrinsic and extrinsic factors such as parent tree age/size, understory light availability and deer browsing, which, under closed canopy conditions are important limiting factors for stump survival or sprout growth. However, the combined effect of potentially confounding abiotic and biotic factors on stump survival and sprout growth remains elusive, even more so under closed canopy conditions. This study aims to quantify the effect of deer browsing on stump mortality and sprout growth under closed-canopy conditions and to compare this effect with other known determinants. Here we show that stump survival and sprout growth in sessile oak (Quercus petraea Matt.) depend on deer browsing, understory light availability and the diameter of the parent tree. By studying paired fenced-unfenced plots, we confirmed that deer browsing decreased stump survival and inhibited sprout growth. Furthermore, by taking advantage of a gradient in understory light availability in monospecific and mixed stands of sessile oak and Scots pine (Pinus sylvestris L.), we showed a clear positive linear relationship between sprout growth and light availability. This relationship explained the observed differences among stand composition types. Finally, we found that increased understory light availability did not compensate for losses due to deer browsing. In the absence of deer browsing, our results demonstrate that sessile oak stumps regenerate well under closed-canopy conditions and maintain a moderate sprouting capacity at least until the age of 70-80 years old. Partial thinning could therefore be a potential tool to renew light-demanding tree species such as sessile oak in mixed high-forest stands. Nevertheless, we do not recommend coppicing sessile oak under closed canopies unless the oak stumps are protected from deer browsing and understory light availability is optimized as much as possible despite closed-canopy-management objectives.

Whereas improved tree mineral nutrition is known to be one driver behind the higher growth in mixed vs pure stands, the effects of stand density and admixture of species with limited functional dissimilarity remain very poorly documented. Our study therefore addresses the impact of local neighbourhood, including density and species composition, on tree nutrition for two species with a limited contrast in traits. We investigated foliar nutrition in a young mixed plantation of beech ( Fagus sylvatica L.) and sycamore maple ( Acer pseudoplatanus L.), planted along a double gradient of density and species proportion. Target trees of both species were selected over the full diameter range along this double gradient, and neighbourhood (local density, species identity and diversity) was determined in a 3 m radius. Using standardized regression analysis, we investigated direct effects of tree size and neighbourhood, and indirect effects of neighbourhood through tree size. Mostly we observed positive tree size effects in both species, be it stronger in beech. For maple, direct positive species identity effects were observed for C, N, P, S, K, Mg, Mn, Zn and Al. For beech, we found indirect negative effects of stand density and diversity. These results were in general agreement with growth data, which suggests that nutrition is one of the driving factors behind complementarity in this stand. We conclude that at this stage the effect of tree size dominates over the neighbourhood effects, including density and diversity. However, the presence of beech reduced intraspecific competition in maple with a positive effect on the foliar nutrient content. As neighbourhood effects are expected to increase as the plantation matures, our findings highlight the potential of admixing tree species with limited dissimilarity in traits for practical use in forestry.

The success of mixed-species tree plantations depends on the balance between positive and negative interactions. Mixtures of Acacia mangium and Eucalyptus urophylla × grandis out-yield their respective monocultures in term of wood production on the Congolese coastal plain, suggesting that facilitation and/or competitive reduction surpass interspecific competition. We investigated how these interactions affected biomass production and N accumulation during the early growth stage of a second rotation of a mixed-species stand of these two species. We used the 15N dilution method to estimate symbiotic nitrogen fixation and its contribution to N accumulation in acacia monoculture and mixture, and we assessed how much N derived from the atmosphere is transferred to the eucalypt trees in the mixed-species stand. Eucalypts grew taller and acacias grew larger in the mixture compared to the monocultures. N mineralomass was greater in the mixture relative to the average values in the two monocultures, with both species contributing to this enhanced N mineralomass. The amount of N derived from the atmosphere in the mixture was 60% higher than that expected given the amount found in acacia monoculture, and 16% of the nitrogen accumulated in eucalypt trees and aboveground eucalypt litterfall was derived from the atmosphere. Reduced competition for light and soil water also contributed to the increased growth of acacias in the mixture, showing that both species benefit from growing in a mixed stand. (Resume d'auteur)

The effects of mixed stands on biodiversity are increasingly being studied since they are supposed to offer higher habitat heterogeneity. Nevertheless, for tree-associated diversity, including epiphytes and terricolous species near tree trunks, few studies exist, and still fewer compare mixed stands with each corresponding pure tree species stand. We evaluated and quantified the influence of forest composition on tree-associated bryophyte diversity (species richness, abundance, composition) in mixed and pure oak-pine stands in a French lowland forest. The main explanatory variables for bryophyte diversity at tree-level were the identity of the phorophyte tree species and the mixture type (pure versus mixed). At the plot level, the main explanatory variable was the stand type (pure oak, pure pine and mixed). We also explored the role of other variables including the chemistry of the bryophyte substrates (soil, bark) and water supply (stemflow, throughfall), as well as stand abundance variables (basal area, interfering plant cover). We analyzed data with Generalized Linear Models under Bayesian statistics, to take into account the spatial autocorrelation between plots and any under- or over-dispersion of our data. At the tree-level, bryophyte richness and abundance were higher on oak than on pine. Pine bryophyte richness was higher in mixed compared to pure stands, whereas for oak, mixed stands did not enhance bryophyte richness. At the plot level, mixed stands hosted bryophyte communities of similar richness to those in pure oak stands, whereas pure pine stands were clearly poorer. Our exploratory models suggested strong effects of water supply chemistry (stemflow and throughfall pH or conductivity) and basal area; the latter had a strong quadratic effect on epiphytic richness at the plot level. In terms of composition, three species were more likely to be found on pine phorophytes, seven species clearly occurred more frequently on oaks. Some species were more likely to be associated to pine in mixed than in pure stands, and one species was found more often under pines in pure stands. Therefore, bryophyte diversity at the landscape level should benefit from the simultaneous presence of the three stand composition types – pure oak, pure pine and mixed stands.

Article résumant et vulgarisant les résultats principaux de la thèse de Maude Toïgo. D'une part, nous exposons l'effet des conditions environnementales sur l'influence du mélange sur la productivité des forêts à partir des données de l'Inventaire Forestier de l'IGN. D'autre part, nous exposons l'effet du mélange sur la largeur et la densité des cernes en interaction avec le déficit en eau dans les peuplements mélangés chêne sessile - pin sylvestre de la forêt d'Orléans.

National audience