Negative Density Dependence Research Articles

An experimental test of the Allee effect range limitation hypothesis.

Understanding how climate change impacts trailing‐edge populations requires information about how abiotic and biotic factors limit their distributions. Theory indicates that socially mediated Allee effects can limit species distributions by suppressing growth rates of peripheral populations when social information is scarce.The goal of our research was to determine if socially mediated Allee effects limit the distribution of Canada warbler Cardellina canadensis at the trailing‐edge of the geographic range.Using 4 years of observational data from 71 sites and experimental data at 10 sites, we tested two predictions of the socially mediated range limitation hypothesis: (a) local growth rates should be positively correlated with local density and (b) the addition of social cues immediately outside the trailing‐edge range boundary would result in colonization of formerly unoccupied habitat and increased growth rates. During the third breeding season, social cues were experimentally added at 10 formerly unoccupied sites within and beyond the species’ local range margin to determine if the addition of social information could increase density and effectively expand the species’ range.No experimental sites were colonized after adding social cues and no evidence of Allee effects was found. Rather, temperature, precipitation and negative density dependence strongly influenced population growth rates.Although theoretical models indicate that the presence of socially mediated Allee effects at species range boundaries could increase the rate of climate‐induced range shifts and local extinctions, empirical results from the first test of this hypothesis suggest that Allee effects play a minimal role in limiting species’ distributions.

Effects of biotic/abiotic factors on the seedling regeneration of Dacrydium pectinatum formations in tropical montane forests on Hainan Island, China

Exploring natural regeneration facilitates a better understanding of the mechanisms of community assembly and biodiversity formation. However, convincing data on tree regeneration in threatened tropical plant communities are lacking. Dacrydium pectinatum is a constructive and endangered species in tropical montane forests of Hainan Island. This study aims to identify the seedling regeneration patterns of D. pectinatum formation and their key drivers. A total of 204 seedling plots were investigated in the northwest (Bawangling, 90 plots), southwest (Jianfengling, 90 plots) and southeast (Diaoluoshan, 24 plots) of Hainan Island. We examined the variations in the seedling/adult composition, density and richness along local scales and elevation gradients and analyzed the relationships between biotic/abiotic variables and seedling regeneration. The seedling distribution pattern changed with environmental gradients, and the seedling density and richness decreased from low to high latitudes and from low to high elevations. Additionally, the adult density and total base area also showed consistent trends. At the community and species levels, the biotic neighborhood variables had a primary impact on the seedling density and richness. Multiple environmental factors (such as the canopy density, elevation, slope, soil organic matter, and soil total nitrogen) also showed significant correlations with the seedling density and richness. Our study supports conspecific negative density dependence and niche filtering as the general mechanisms of natural regeneration of D. pectinatum formations. Both biotic and abiotic factors might affect the survival and growth of seedlings, and their relative importance varies with environmental gradients and species. We conclude that enclosing forests with moderate stand densities in low-elevation areas to reduce human interference will help optimize the community structure. However, for high-elevation areas and high-density forests, appropriate thinning, manual regeneration and open gaps are needed to improve the light environment and soil nutrients, thereby promoting the natural regeneration of D. pectinatum formations.

Density dependence and habitat heterogeneity regulate seedling survival in a North American temperate forest

Survival and growth of forest tree seedlings are influenced by many abiotic and biotic forces that may vary across space and time. The simultaneous influences of habitat heterogeneity, temporal environmental variability (including disturbance regimes), and biotic interactions are difficult to disentangle, yet understanding the relative importance of these factors for tree seedling dynamics is critical for conservation of forest biodiversity. Most long term, spatially explicit studies of tree seedling ecology have been set in the tropics; much less attention has been given to tree seedlings in temperate forests. We monitored the survival of over 3000 individual seedlings over an eight-year period in a temperate forest in northeast Wisconsin, USA. Results from four seedling censuses demonstrated that both conspecific density and environmental variables significantly affected seedling survival. Higher densities of conspecific trees consistently reduced the probability of seedling survival over time. At the community level, relatively common species were negatively influenced by neighboring conspecific trees and exhibited higher per capita and per basal area mortality. The negative impacts of high conspecific tree density were most pronounced in areas of higher light and moisture, but these interactions varied over time, as did the importance of other abiotic variables. Patterns of less common species were more clearly explained by abiotic variables, with shifting relevance of specific variables according to species abundance. Conspecific negative density dependence, which interacts with resource gradients, and habitat conditions that shift over time are influencing community composition in this northern temperate forest.

Environmental conditions modulate compensatory effects of site dependence in a food-caching passerine.

Although density regulates the abundance of most wild animal populations by influencing vital rates, such as fecundity and survival, the mechanisms responsible for generating negative density dependence are unclear for many species. Site dependence occurs when there is preferential filling of high-quality territories, which results in higher per capita vital rates at low densities because a larger proportion of occupied territories are of high quality. Using 41yr of territory occupancy and demographic data, we investigated whether site dependence was a mechanism acting to influence fecundity and, by extension, regulate a population of Canada Jays in Algonquin Provincial Park, Ontario, Canada. As predicted by site dependence, the proportion of occupied territories that were of high quality was negatively correlated with population density and periods of vacancy were shorter for high-quality territories than for low-quality territories. We also found evidence that per capita fecundity was positively related to the proportion of occupied territories that were of high quality, but only when environmental conditions, which influence the entire population, were otherwise poor for breeding. Our results suggest that site dependence likely plays a role in regulating this population but that environmental conditions can modulate the strength of density dependence.

Seedling survival declines with increasing conspecific density in a common temperate tree

AbstractFeedbacks between plants and their soil microbial communities often drive negative density dependence in rare, tropical tree species, but their importance to common, temperate trees remains unclear. Additionally, whether negative density dependence is driven by natural enemies (e.g., soil pathogens) or by high densities of seedlings has rarely been assessed. Density dependence may also depend on seedling size, as smaller and/or younger seedlings may be more susceptible to mortality agents. We monitored seedlings of Quercus rubra, a common, canopy‐dominant temperate tree, to investigate how the density of neighboring adults and seedlings influenced their survival over two years. We assessed how the soil microbial community influenced seedling survival by growing seedlings in a glasshouse inoculated with soil collected from beneath conspecific and heterospecific mature trees. In the field, seedling survival was lower in areas with high densities of mature conspecifics but was unrelated to either conspecific or heterospecific seedling density. Smaller seedlings were also more sensitive than larger seedlings to neighboring adult conspecifics. In the glasshouse, seedlings grown with soil from beneath a conspecific adult had a higher mortality rate than seedlings grown with soil from beneath heterospecific adults or sterilized soil, suggesting that soil microbial communities drive the patterns of mortality in the field. These results illustrate the importance of negative density‐dependent feedbacks resulting from the soil microbial community in a common and ecologically important temperate tree species.

The effects of density dependence and habitat preference on species coexistence and relative abundance.

In plant communities, some mechanisms maintain differences in species' abundances, while other mechanisms promote coexistence. Asymmetry in conspecific negative density dependence (CNDD) and/or habitat preference is hypothesized to shape relative species abundance, whereas community compensatory trends (CCTs) induced by community-level CNDD and heterospecific facilitation are hypothesized to promote coexistence. We use survey data from three 1-ha permanent dynamic plots in a subtropical forest over the course of a decade to find out which of these processes are important and at which life-history stages (the seedling, sapling, and juvenile stages) they exert their effects. CNDD was not related to abundance in any of the life-history stages. Suitable habitats positively influenced plant abundance at all tested life stages, but especially so for juveniles. Community-level CNDD of seedling neighbors was detected at the seedling stage, while heterospecific facilitation was detected across all tested life-history stages. A CCT in seedling survival was detected, but there was no evidence for such trends across the other life-history stages. Altogether, our results suggest that habitat specificity increases the rarity of species, whereas a CCT at the seedling stage, which is likely to be induced by CNDD and heterospecific facilitation, enables such species to maintain their populations.

Negative density-dependence of the structural relaxation time of liquid silica: insights from a comparative molecular dynamics study

In many tetrahedral network-forming liquids, structural relaxation is anomalously accelerated by compression over relatively low pressure ranges. Here, for silica, we study this problem through comparative molecular dynamics simulations using two different models. Under compression, the network structures are compacted by slight tuning of the intertetrahedral bond angles while nearly preserving the unit tetrahedral structure. The consequent structural changes are remarkable for length scales larger than the nearest neighbor ion-pair distances. Accompanying with such structural changes, the interactions of the nearest Si–O pairs remain almost unchanged, whereas those of other ion pairs are, on average, strengthened by the degree of compression. In particular, the enhancement of the net Si–O interactions at the next nearest neighbor distance, which assist an ion in escaping from the potential well, reduces the activation energy, leading to a significant acceleration of structural relaxation. The results of our comparative molecular dynamics simulations are compatible with the scenario proposed by Angell, and further indicate that the structural relaxation dynamics cannot be uniquely determined by the configurations but strongly depends on the details of the coupling between the structure and the interaction.

Soil biota increase the likelihood for coexistence among competing plant species.

Theory predicts that stable species coexistence will occur when population growth rates of competitively dominant species are suppressed when at high conspecific density. Although there is now compelling evidence that plant communities exhibit negative density dependence, the relative importance of the underlying processes leading to these patterns is rarely tested. We coupled reciprocal greenhouse and field experiments with community dynamics modeling to untangle the relative importance of soil biota from competition as stabilizing forces to coexistence. We found that (1) plant-soil biotic interactions compared to competitive interactions were stronger stabilizing forces, (2) only the strength of plant-soil biotic interactions was dependent on plant evolutionary history, and (3) the variation in the strength of plant-soil biotic interactions was correlated with relative abundance patterns in an opposite way than was the variation in the strength of competitive interactions. Collectively, our results demonstrate the fundamental role soil biota have in maintaining plant community diversity.

Traits mediate a trade‐off in seedling growth response to light and conspecific density in a diverse subtropical forest

Abstract Understanding tree species responses to biotic and abiotic factors is fundamental for stronger predictions of community assembly and dynamics. However, several challenges remain. These include a failure to investigate whether there is evidence for key hypothesized life‐history trade‐offs and to link these trade‐offs to functional traits. In this study, we seek to explicitly address the above outstanding challenges by constructing models for individual seedling growth in response to abiotic and biotic factors using 3 years of seedling census data from a 20‐ha subtropical forest dynamics plot in a diverse subtropical forest and correlated these responses with functional traits. We found that light and conspecific neighbours increase and decrease the RGR of tree seedlings respectively. We also found that the ability of a species to positively respond to canopy openness trades‐off against susceptibility to conspecific negative density dependence (CNDD). This trade‐off was evident across seasons and could be predicted on functional trait—stem and leaf dry matter content. Synthesis. Our findings indicate species that can grow quickly in high‐light environments also tend to suffer more CNDD. The results highlight that strong evidence of a trade‐off relating to growth and defence widely hypothesized to be of importance in diverse tree communities and that this trade‐off occurs across seasons and can be linked to a commonly measured functional trait.

Predation yields greater population performance: What are the contributions of density- and trait-mediated effects?

1. Population responses to extrinsic mortality can yield no change in number of survivors (compensation) or an increase in number of survivors (overcompensation) when the population is regulated by negative density-dependence. This intriguing response has been the subject of theoretical studies, but few experiments have explored how the source of extrinsic mortality affects the response. 2. This study tests abilities of three functionally diverse predators, alone and combined, to induce (over)compensation of a prey population. Larval Aedes aegypti (Diptera: Culicidae) were exposed to predation by Mesocyclops longisetus (Crustacea: Copepoda), Anopheles barberi (Diptera: Culicidae), Corethrella appendiculata (Diptera: Corethrellidae), all three in a substitutive design, or no predation. 3. The number of survivors to adulthood, female size and development time, and a composite index of performance (r') were analysed. Predator treatment did not have a significant effect on total number of survivors, nor on number of males, suggesting mortality by predation was compensatory. Predation significantly affected number of female survivors, with a trend of more females produced with predation, though no post hoc tests were significant. Predation significantly increased female development rate and r' relative to no-predator control. 4. A sensitivity analysis indicated that the change in the number of female adults produced was the largest contributing factor to the differences in r' among cohorts. While predation did not significantly increase overall production of adults, it did release survivors from density-dependent effects sufficiently to increase population performance. This study provides an empirical test of mechanisms by which predation may yield positive effects on a population of victims, a phenomenon predicted to occur across many taxa and food webs.

On the need to evaluate costs and benefits of synzoochory for plant populations

Abstract Many species interactions occur along a continuum from mutualistic to antagonistic. Among these interactions, the effects of seed‐dispersing granivores (synzoochory) are especially interesting because the benefits and costs are so striking. Here, we respond to a commentary by C. M. Moore and J. W. Dittel (‘On mutualism, models, and masting: the effects of seed‐dispersing animals on the plants they disperse’). We focus on the points that were most thought‐provoking to us, specifically the notions that (a) synzoochory is widespread, and therefore ought to be beneficial for plants, (b) dispersal should reduce negative density dependence and (c) plant life spans are at least an order of magnitude longer than typical study lengths. We argue that (a) widespread occurrence of an interaction cannot serve as an argument that this interaction is mutualistic, (b) dispersal effects on density dependence are variable and need to be tested for every particular species pair and (c) Short‐term studies can be used to gain mechanistic understanding of interactions even without risky long‐term extrapolations. Synthesis. Evaluating the costs and benefits of synzoochory for plant populations under different ecological contexts will help to understand the ecological principles of the interaction but also allows to move the interaction towards each end of the antagonism–mutualism continuum in nature management and conservation.

Conspecific negative density dependence in rainy season enhanced seedling diversity across habitats in a tropical forest.

Conspecific negative density dependence (CNDD) could be one of the most important local-scale mechanisms shaping plant species coexistence. However, the spatial and temporal changes in the strength CNDD and the implications for the plant diversity remain unknown. We used 10years of seedling data, in a seasonal tropical rainforest, to discover how CNDD influences tree seedling survival across habitats and seasons. We also evaluated the relation between CNDD and species diversity. We found the strength of CNDD in the valley habitat was significantly stronger than in ridge habitat in rainy season, but not significantly different in dry season. Corresponding to expectations of CNDD as mechanism of diversity maintenance, seedling species diversity was significantly higher in valley habitat than in ridge habitat and significantly correlated with CNDD. Additionally, conspecific and heterospecific seedling neighbour densities positively affected the survival of tree seedlings, but heterospecific adult neighbour density had a weak effect. Our study finds that CNDD varied significantly across habitats and was correlated with local seedling diversity. Our results highlight the importance of CNDD in driving species diversity at the local scale. Recognizing the spatial and temporal variation in the strength of CNDD will aid efforts to model and understand species coexistence.

Surprising low diversity of the plant pathogen Phytophthora in Amazonian forests.

The genus Phytophthora represents a group of plant pathogens with broad global distribution. The majority of them cause the collar and root-rot of diverse plant species. Little is known about Phytophthora communities in forest ecosystems, especially in the Neotropical forests where natural enemies could maintain the huge plant diversity via negative density dependence. We characterized the diversity of soil-borne Phytophthora communities in the North French Guiana rainforest and investigated how they are structured by host identity and environmental factors. In this little-explored habitat, 250 soil cores were sampled from 10 plots hosting 10 different plant families across three forest environments (Terra Firme, Seasonally Flooded and White Sand). Phytophthora diversity was studied using a baiting approach and metabarcoding (High-Throughput Sequencing) on environmental DNA extracted from both soil samples and baiting-leaves. These three approaches revealed very similar communities, characterized by an unexpected low diversity of Phytophthora species, with the dominance of two cryptic species close to Phytophthora heveae. As expected, the Phytophthora community composition of the French Guiana rainforest was significantly impacted by the host plant family and environment. However, these plant pathogen communities are very small and are dominated by generalist species, questioning their potential roles as drivers of plant diversity in these Amazonian forests.

Positive density dependence acting on mortality can help maintain species-rich communities.

Conspecific negative density dependence is ubiquitous and has long been recognized as an important factor favoring the coexistence of competing species at local scale. By contrast, a positive density-dependent growth rate is thought to favor species exclusion by inhibiting the growth of less competitive species. Yet, such conspecific positive density dependence often reduces extrinsic mortality (e.g. reduced predation), which favors species exclusion in the first place. Here, using a combination of analytical derivations and numerical simulations, I show that this form of positive density dependence can favor the existence of equilibrium points characterized by species coexistence. Those equilibria are not globally stable, but allow the maintenance of species-rich communities in multispecies simulations. Therefore, conspecific positive density dependence does not necessarily favor species exclusion. On the contrary, some forms of conspecific positive density dependence may even help maintain species richness in natural communities. These results should stimulate further investigations into the precise mechanisms underlying density dependence.

Establishment of a hemiparasite Rhinanthus alectorolophus and its density‐dependent suppressing effect on a grass: A case study from golf roughs

AbstractSuppression effect of hemiparasites on grasses is a phenomenon, that can be utilized to increase biodiversity of various grassland systems. Host suitability, environmental effects, and hemiparasite abundance most likely influence the impact of parasitism on host height and biomass. However, the role of these factors is unclear in field conditions. We studied host suitability, establishment rates, and effects of a hemiparasitic plant Greater yellow rattle [ Rhinanthus alectorolophus (Scop.)] (RA) on canopy height and biomass of a grass tall fescue (Festuca arundinacea Schreb.) in five trials with manipulated fertilization conducted in golf roughs. Particularly, we examined how grass height and biomass changed with increasing hemiparasite abundance. On average, grasses (nine in total) were better hosts than legumes (four in total). Tall fescue belonged to the group of better hosts. The average establishment rate of Rhinanthus plants ranged from 4.4 to 24.8% regardless of fertilization treatments. In fertilization trials, Rhinanthus did not affect grass height but weakly suppressed grass biomass in non‐fertilized plots. The relationship between grass height/biomass and Rhinanthus abundance was best described by a logarithmic decay model. The suppressive effect tended to saturate (negative density dependence) in plots where the Rhinanthus abundance was very high. However, such high densities were quite rare and low Rhinanthus abundances were usually insufficient to substantially suppress grass growth. Our results indicate that we should aim for at least 50 Rhinanthus plants per 1 m2 to achieve desirable effects on biodiversity and that Rhinanthus performance will be better when nutrients are rather limiting.

Trait-mediated neighbor effects on plant survival depend on life stages and stage-specific traits in a temperate forest

Among the multiple ecological processes that determine species coexistence and community dynamics, conspecific negative density dependence (CNDD) and niche partitioning are perhaps among the most important. However, the relative strength of these mechanisms may change across life stages and depend on species ecological strategies. We used a dataset that includes information on stage-specific functional traits and a long-term, spatially explicit data of plant survival including 7971 individuals and 41 species monitored for seedling and adult life stages to examine the relative role of these mechanisms influencing individual survival across ontogenetic stages in a 9-ha temperate forest plot in China. Our results showed that for seedlings, CNDD became weaker for species with larger leaf area, while for adults, the conspecific neighbors had a positive effect on survival and this effect became stronger with decreasing seed mass. Also, the results examing the effects of neighboring heterospecifics suggest a prevalence of niche partitioning at the seedling stage but a prevalence of environmental filtering or hierarchical competition for the adult stage. In particular, the strength of these processes at the adult life stage increased with higher maximum height and smaller specific leaf area. Finally, we found that the effect of traits on individual survival for seedling and adult life stages was dependent on both the prevalence of conspecifics and functional diversity of heterospecifics in the neighborhood. Overall, our results demonstrates that ecological processes tend to have a stronger diversifying effect for seedlings than for adults and these effects depend on stage-specific species traits. Considering trait variation and their interacting effects with biotic factors across life stages provides further insights into ecological mechanisms shaping plant communities and driving its dynamics.

Tree mycorrhizal type mediates the strength of negative density dependence in temperate forests

Abstract Recent plant–soil feedback experiments suggest that arbuscular mycorrhizal (AM) tree species experience stronger conspecific negative density dependence (CNDD) than ectomycorrhizal (EM) tree species. Yet, how these findings inform our understanding of natural systems is limited because the roles of local soil conditions, light availability and tree species abundances in influencing CNDD for AM and EM species are not clear. Here we examined seedling and sapling survival in two temperate old‐growth forests (broad‐leaved pine and spruce‐fir forests) in Northeast China, to evaluate the effects of both conspecific and heterospecific neighbour density, as well as the soil and light environments, on the survival of AM‐ and EM‐dependent trees at early life stages. While light availability increased the survival of EM seedlings, soil organic resources increased EM sapling survival in the spruce‐fir plot. AM tree species suffered stronger CNDD than did EM tree species in both plots. In the spruce‐fir plot, soil factors and light availability mediated species CNDD but their effects differed between AM and EM species, and also between seedlings and saplings. For seedlings in both plots, we found that AM species exhibited a positive relationship between species abundance and CNDD strength, whereas this relationship was negative for EM species. Synthesis. Our results provide one of the few tests of how fungal symbioses determine species responses to intraspecific and interspecific interactions and the direct effects of local environmental conditions on seedling and sapling survival. We show that mycorrhizal type mediates the strength of CNDD and its relationship with species abundance. These results suggest that tree mycorrhizal association can determine the strength of CNDD effect on both rare and common species, and these CNDD differences are likely to influence the community composition of temperate forests.

The asymmetric relationships of the distribution of conspecific saplings and adults in forest fragments

Abstract Aims With the increase of land-use change, habitat fragmentation has become a major factor affecting plant diversity. Generally, when the adult density is high, the survival rate of conspecific saplings may decline and provide more resources and space for other species to maintain a high local species diversity. Therefore, the spatial relationship of conspecific saplings and adults (SRCSA) can regulate plant diversity. However, very few studies have assessed SRCSA within fragmented forests, and we still largely unknown the relationship between SRCSA and species diversity in fragmented landscapes. Methods We calculated the mean strength of SRCSA using the spatial distribution data of saplings and adults with a log-transformed hierarchical offset-power model on 27 islands in a land-bridge island system. The higher strength of SRCSA reflects a more negative relationship between the density of conspecific saplings and adults on an island (i.e. the looser pattern of SRCSA). We tested the relationships among island attributes (i.e. island area, the distance to the mainland—MD, and the distance to the nearest island—ND), SRCSA and species diversity. Important Findings The strength of SRCSA increased with ND. Meanwhile, the species diversity significantly increased with the strength of SRCSA, and island area and the strength of SRCSA independently explained 26% and 6% of variation of species diversity, respectively, and their interactions explained 8%. Shade-intolerant and low-abundant species showed looser patterns of SRCSA. Our study suggests that SRCSA may have the effect of excluding species with a relatively low abundance in isolated island forests, which illustrates the importance of biotic interactions in maintaining plant diversity in fragmented forests. Our results also emphasize that we should consider forest connectivity when testing the conspecific negative density dependence.

Mutualist and pathogen traits interact to affect plant community structure in a spatially explicit model

Empirical studies show that plant-soil feedbacks (PSF) can generate negative density dependent (NDD) recruitment capable of maintaining plant community diversity at landscape scales. However, the observation that common plants often exhibit relatively weaker NDD than rare plants at local scales is difficult to reconcile with the maintenance of overall plant diversity. We develop a spatially explicit simulation model that tracks the community dynamics of microbial mutualists, pathogens, and their plant hosts. We find that net PSF effects vary as a function of both host abundance and key microbial traits (e.g., host affinity) in ways that are compatible with both common plants exhibiting relatively weaker local NDD, while promoting overall species diversity. The model generates a series of testable predictions linking key microbial traits and the relative abundance of host species, to the strength and scale of PSF and overall plant community diversity.

Variations of density-dependent seedling survival in a temperate forest

Seedling dynamics are driven by various biotic factors (for example, neighborhood interactions) and abiotic conditions (for example, light availability, topography, and soil nutrients). Therefore, quantifying the drivers of seedling survival is critical for understanding seedling dynamics within plant communities. However, how the importance of certain biotic and abiotic factors changes over growth stages is still relatively unknown. In the present study, the importance of density and distance dependence were examined, as well as the effects of abiotic factors, in regard to the survival of both newly germinated and established seedlings and saplings (<1 cm DBH; taller than 30 cm in height; and at least 2 years of age). A total of 1305 seedlings belonging to 21 species of a temperate forest in northeastern China were monitored. It was found there were significant conspecific negative density dependence (CNDD) effects impacting the survival of the established seedlings. In addition, the newly germinated seedlings were found to be less affected by the CNDD. The abiotic effects on seedling survival were not found to be significant. These results suggested that at the community level, the density dependence effects may be stronger during the seedling stage when compared with the abiotic factors. The distance dependence was observed to be mainly effective within a radius of approximately 5 m for both the newly germinated and established seedling communities. The directions and magnitudes of the aforementioned factors were affected by different life-history strategies. The seedlings which were wind-dispersed and the shade-tolerant species tended to suffer less from the CNDD. Therefore, based on an unusually large set of observations in a temperate forest, this study identified and analyzed the factors affecting seedling survival for different life-history strategies. The obtained results will potentially provide new perspectives for deepening the understanding of the dynamics of temperate forests.