Tree Recruitment Research Articles

Species-specific spatial associations shape tree recruitment dynamics in an arid mixed forest of Picea crassifolia and Pinus tabuliformis

Species-specific spatial associations shape tree recruitment dynamics in an arid mixed forest of Picea crassifolia and Pinus tabuliformis

The need for carbon finance schemes to tackle overexploitation of tropical forest wildlife.

Defaunation of tropical forests, particularly from unsustainable hunting, has diminished populations of key seed dispersers for many tree species, driving shifts in forest community composition toward small-fruited or wind-dispersed trees with low wood density. Such shifts can reduce aboveground biomass, prompting calls for overexploitation to be included in bioeconomic policy, but a synthesis of existing literature for wildlife impacts on carbon stores is lacking. We evaluated the role of wildlife in tropical forest tree recruitment and found that it was critical to tropical forest carbon dynamics. The emerging financial value of ecosystem services provided by tropical forest fauna highlights the need for carbon-based payments for ecosystem services schemes to include wildlife protection. We argue for three cost-effective actions within carbon finance schemes that can facilitate wildlife protection: support land security opportunities for Indigenous peoples and local communities; provide support for local people to protect forest fauna from overexploitation; and focus on natural regeneration in restoration projects. Incorporating defaunation in carbon-financing schemes more broadly requires an increased duration of carbon projects and an improved understanding of defaunation impacts on carbon stores and ecosystem-level models. Without urgent action to halt wildlife losses and prevent empty forest syndrome, the crucial role of tropical forests in tackling climate change may be in jeopardy.

The phenology of Eucalyptus camaldulensis (Dehnh, 1832) and Eucalyptus coolabah (Blakely & Jacobs, 1934) in the northern Murray–Darling Basin and implications for recruitment on floodplains

Context Published evidence suggests recruitment of floodplain trees is declining in the northern Murray–Darling Basin (MDB) and that this is linked to river flow. Aims We investigated the reproductive phenology of Eucalyptus camaldulensis and Eucalyptus coolabah to better understand tree reproductive cycles and the influence of rainfall and flow on seed production. Methods We surveyed phenological events (bud, flower and fruit production) over 6 years, at nine sites along the Condamine–Balonne River and examined correlations between these events and meteorological parameters. Key results For both species, event timing was consistent across sites and years and event relative abundance scores were different between years. Bud abundance was correlated with rainfall, flooding and warming temperatures and E. coolabah produced additional bud crops after floods. Conclusions The timing of events did not differ significantly from published results for the southern MDB. Rainfall, river flows and flooding in spring resulted in more abundant bud crops in E. camaldulensis and E. coolabah, and subsequently more abundant fruit crops. Fruit dehiscence coincided with summer–autumn flooding, which provides seed for germination. Genetic and physiological factors may also influence the observed pattern. Implications These results will inform water planning in Queensland, Commonwealth Environmental Water Holder watering actions and long-term watering plans in the northern MDB.

Demographic and net primary productivity dynamics of primary and secondary tropical forests in Southwest China under a changing climate

AbstractTropical forests are major carbon sinks on the Earth's land surface. However, our understanding of how the demographic rate and carbon sink capacities of tropical forests respond to climate change remains limited. In this study, we investigated the impacts of environmental drivers on forest growth, mortality, recruitment, and stem net primary productivity (NPPstem) over 16 years at five tropical forest plots in Xishuangbanna, Southwest China. These plots are along a successional gradient spanning three tropical secondary forests (tropical secondary forest‐1 [TSF‐1], tropical secondary forest‐2 [TSF‐2], and tropical secondary forest‐3 [TSF‐3]) and two primary forests (tropical rainforest [TRF] and tropical karst forest [TKF]). Our results showed that early successional secondary forests (TSF‐2 and TSF‐3) had higher diameter growth rates and relative mortality rates. An extreme drought event during 2009–2010 reduced the growth rate, relative recruitment rate, and NPPstem for most plots while increasing mortality in early successional forest plots. We observed significant negative effects of maximum temperature (Tmax) on NPPstem and diameter growth rate across all plots. Additionally, we found that precipitation had significant positive effects on diameter growth rate across all plots. Furthermore, tree mortality increased with rising Tmax, whereas precipitation significantly enhanced tree recruitment. Our findings highlight the vulnerability of tree growth, mortality, recruitment, and productivity in tropical forests to extreme drought events in Southwest China. Continued climate warming and more frequent droughts will induce higher mortality rates and impede growth, thus reducing the carbon sink capacity of tropical forests, especially in early successional stage tropical secondary forests.

Understanding woody plant encroachment: A plant functional trait approach

AbstractThe increasing density of woody plants threatens the integrity of grassy ecosystems. It remains unclear if such encroachment can be explained mostly by direct effects of resources on woody plant growth or by indirect effects of disturbances imposing tree recruitment limitation. Here, we investigate whether woody plant functional traits provide a mechanistic understanding of the complex relationships between these resource and disturbance effects. We first assess the role of rainfall, soil fertility, texture, and geomorphology to explain variation in woody plant encroachment (WPE) following livestock grazing and consequent fire suppression across the Serengeti ecosystem. Second, we explore trait‐environment relationships and how these mediate vegetation response to fire suppression. We find that WPE is strongest in areas with high soil fertility, high rainfall, and intermediate catena positions. These conditions also promote woody plant communities characterized by small stature and seed sizes smaller relative to a comparative baseline within the Serengeti ecosystem, alongside high recruit densities (linked to a recruitment‐stature trade‐off). The positioning of species along this “recruitment‐stature axis” predicted woody stem density increase in livestock sites. Structural equation modeling suggested a causal pathway where environmental factors shape the community trait composition, subsequently influencing woody recruit numbers. These numbers, in turn, predicted an area's vulnerability to WPE. Our study underscores the importance of trait‐environment relationships in predicting the impact of human alterations on local vegetation change. Understanding how environmental factors directly (resources) and indirectly (legacy effects and plant traits) determine WPE supports the development of process‐based ecosystem structure and function models.

Recruitment and Aggregation Capacity of Tea Trees to Rhizosphere Soil Characteristic Bacteria Affects the Quality of Tea Leaves.

There are obvious differences in quality between different varieties of the same plant, and it is not clear whether they can be effectively distinguished from each other from a bacterial point of view. In this study, 44 tea tree varieties (Camellia sinensis) were used to analyze the rhizosphere soil bacterial community using high-throughput sequencing technology, and five types of machine deep learning were used for modeling to obtain characteristic microorganisms that can effectively differentiate different varieties, and validation was performed. The relationship between characteristic microorganisms, soil nutrient transformation, and tea quality formation was further analyzed. It was found that 44 tea tree varieties were classified into two groups (group A and group B) and the characteristic bacteria that distinguished them came from 23 genera. Secondly, the content of rhizosphere soil available nutrients (available nitrogen, available phosphorus, and available potassium) and tea quality indexes (tea polyphenols, theanine, and caffeine) was significantly higher in group A than in group B. The classification result based on both was consistent with the above bacteria. This study provides a new insight and research methodology into the main reasons for the formation of quality differences among different varieties of the same plant.

Dynamics of tree stems and biomass in old‐growth and secondary forests along gradients in liana dominance, elevation and soil

Abstract Lianas, or woody vines, are key components of many tropical forests and can have substantial impacts on the dynamics and functioning of these important ecosystems. Their competition with trees for resources, in particular light, can hamper the recovery of forests from disturbances. Yet, it is unclear how forest disturbance interacts with liana–tree ratio (LTR), topography and soil properties to shape tree dynamics and the trajectories of forest succession. Using temporal data from the Kilombero Valley and the Udzungwa Mountains of Tanzania, we demonstrate how the dynamics of tree stems and biomass vary between secondary and old‐growth forests with changes in the dominance of lianas and environmental gradients. Greater tree recruitment and mortality in secondary forests compared with old‐growth forests suggested rapid regeneration processes and faster turnover. However, no significant differences were found in the net annual changes in the number or biomass of trees between secondary and old‐growth forests. Our findings also showed that higher LTRs were positively associated with stem mortality but also with tree biomass growth, indicating a nuanced ecological role of lianas in forest ecosystems, which warrants further investigation to fully understand the causal factors at play. Net changes in tree stem numbers decreased significantly with elevation, implying climatic constraints on forest regeneration at higher elevations. Soil cation exchange capacity and organic carbon were found to significantly influence tree stem recruitment and net change in abundance, although their effects on biomass remained unclear. Synthesis: Our findings indicate that the recovery of tropical forests from disturbance in terms of the number and biomass of tree stems may be predictable along environmental gradients. These insights have the potential to broaden our capacity to develop more nuanced strategies that identify when and where tropical forests may require restoration interventions, with a focus on structural recovery.

Grow or die: A 49‐year growth history of a Japanese warm‐temperate tree species

AbstractThe growth trajectories of trees are not fully understood due to their long life span. We characterized the population dynamics of the canopy tree Castanopsis cuspidata (Thunb.) Schottky in a Japanese warm‐temperate forest over 49 years (1966–2015). Our study was initiated approximately 50 years after our study site was clear‐cut. The forest had a closed canopy for the first 23 years of the study. Strong typhoons in 1991 and 1993 seriously damaged the forest, and since then the forest has been recovering from these disturbances. The diameter distribution of this species was bell‐shaped in 1966, suggesting that the trees emerged simultaneously after the clear‐cut in the 1910s, and the recruitment of trees has remained unchanged since then. The lack of recruitment of C. cuspidata before the typhoon disturbance supports this conclusion. Assuming that the C. cuspidata trees in 1966 were cohorts that were established soon after the clear‐cut, the size differences reflect differences in growth rate, with small trees corresponding to slow growers and large trees corresponding to fast growers. Before the typhoon, slow growers had low survival, and the mortality rate of fast growers was low. Many fast growers were uprooted or snapped by strong winds by the typhoons. However, their mortality rate did not differ from that of slow growers because many slow growers were killed by large fallen trees. The growth of some slow‐growing survivors increased after the typhoon, which allowed them to rapidly reach the canopy. Therefore, the typhoon altered the distribution of canopy trees among slow and fast growers. Survivors experienced faster growth than trees that died during the census period, suggesting that growth rate provides a robust indicator of future survival. Before the typhoon, the survival of fast growers was higher than that of slow growers. This suggests that fast growers disproportionally contribute to reproduction compared with slow growers. However, no recruited tree was observed in this subperiod, suggesting that fast growers made no contribution to reproduction. Fast growers might not play a more significant demographic role than slow growers in this species.

Facilitation drives tree seedling survival at alpine treelines

Abstract Alpine treelines are considered ecological monitors recording the impacts of climate change on trees and forests. To date, most treeline research has focused on how climate change drives treeline dynamics. However, little is known about how biotic interactions mediate treeline shifts, particularly in the case of tree recruitment, a bottleneck of treeline dynamics. We hypothesized that inter- and intraspecific facilitation determined the establishment and survival of tree seedlings at alpine treelines. To test this hypothesis, 630 Abies georgei var. smithii seedlings with different ages (4–6, 7–9 and 10–15 years old) were transplanted into three growth habitats (canopy-in, canopy-out and meadow) across the alpine treeline ecotone (4300–4500 m) in the Sygera Mountains, on the southeastern Qinghai-Tibetan Plateau. Microclimate, height growth, mortality rates and leaf functional traits of transplanted seedlings were measured over 3 years. We found that the variations in leaf functional traits were driven by microclimate. After the transplantation, the leaf concentrations of soluble sugars and starch and C:P ratio increased, whereas leaf size decreased. The resource use of seedlings gradually shifted to a more conservative strategy as indicated by changes in non-structural carbohydrates and nutrient concentrations. Radiation, temperature and moisture conditions, mediated by plant interactions, influenced seedling mortality and annual growth by affecting leaf morphological traits. Our findings illustrate how facilitation plays a crucial role in altering solar radiation and leaf trait functioning, determining seedling survival and growth at alpine treelines. We provide new insights into the underlying mechanisms for tree establishment and alpine treeline shifts in response to climate change.

The puzzling ecology of African Marantaceae forests.

Marantaceae forests are tropical rainforests characterized by a continuous understory layer of perennial giant herbs and a near absence of tree regeneration. Although widespread in West-Central Africa, Marantaceae forests have rarely been considered in the international literature. Yet, they pose key challenges and opportunities for theoretical ecology that transcend the borders of the continent. Specifically, we ask in this review whether open Marantaceae forests and dense closed-canopy forests can be considered as one of the few documented examples of alternative stable states in tropical forests. First, we introduce the different ecological factors that have been posited to drive Marantaceae forests (climate, soil, historical and recent anthropogenic pressures, herbivores) and develop the different hypotheses that have been suggested to explain how Marantaceae forests establish in relation with other vegetation types (understory invasion, early succession after disturbance, and intermediate successional stage). Then, we review the underlying ecological mechanisms that can explain the stability of Marantaceae forests in the long term (tree recruitment inhibition, promotion of and resilience to fire, adaptive reproduction, maintenance by megaherbivores). Although some uncertainties remain and call for further empirical and theoretical research, we found converging evidence that Marantaceae forests are associated with an ecological succession that has been deflected or arrested. If verified, Marantaceae forests may provide a useful model to understand critical transitions in forest ecosystems, which is of particular relevance to achieve sustainable forest management and mitigate global climate change.

Tree regeneration in models of forest dynamics: A key priority for further research

AbstractTree regeneration is a key process in forest dynamics, particularly in the context of forest resilience and climate change. Models are pivotal for assessing long‐term forest dynamics, and they have been in use for more than 50 years. However, there is a need to evaluate their capacity to accurately represent tree regeneration. We assess how well current models capture the overall abundance, species composition, and mortality of tree regeneration. Using 15 models built to capture long‐term forest dynamics at the stand, landscape, and global levels, we simulate tree regeneration at 200 sites representing large environmental gradients across Central Europe. The results are evaluated against extensive data from unmanaged forests. Most of the models overestimate recruitment levels, which is compensated only in some models by high simulated mortality rates in the early stages of individual‐tree dynamics. Simulated species diversity of recruitment generally matches observed ranges. Models simulating higher stand‐level species diversity do not feature higher species diversity in the recruitment layer. The effect of light availability on recruitment levels is captured better than the effects of temperature and soil moisture, but patterns are not consistent across models. Increasing complexity in the tree regeneration modules is not related to higher accuracy of simulated tree recruitment. Furthermore, individual model design is more important than scale (stand, landscape, and global) and approach (empirical and process‐based) for accurately capturing tree regeneration. Despite the mismatches between simulation results and data, it is remarkable that most models capture the essential features of the highly complex process of tree regeneration, while not having been parameterized with such data. We conclude that much can be gained by evaluating and refining the modeling of tree regeneration processes. This has the potential to render long‐term projections of forest dynamics under changing environmental conditions much more robust.

Long-term effects of a tornado: Impacts on woody native vegetation and invasive Amur honeysuckle (Lonicera maackii) in an urban forest.

As tornados become increasingly common with global climate change, recovery of the woody vegetation in temperate forests is imperative to maintain an intact ecosystem. In many urbanized landscapes, invasive species are also increasing and could interfere with natural recovery from environmental disturbance. We quantified the impact and 17-year recovery from a major tornado in a temperate deciduous forest. We used vegetational surveys in southwestern Ohio at the Harris M. Benedict Nature Preserve, where approximately a third of this site was damaged by a tornado in 1999. Plots were established in the tornado-damaged area and the nearby undisturbed forest to examine forest recovery of trees/saplings, shrubs and vines, and tree seedlings during 2003, 2006, 2010, and 2016/2017. The number of tree saplings, shrubs, and vines increased immediately after the tornado, but then declined by 2010, relative to the undisturbed forest. Forest tree recruitment was lower in tornado-damaged sites with fewer tree seedlings, but more saplings. Tree diversity was also affected by Agrilus planipennis (Emerald Ash borer) which targeted native ash trees within this time period. Despite an initial increase in shrubs and vines in the damaged area, the diversity and density of shrubs approached equality in both sites by 2016. Most shrubs in both sites were the invasive Lonicera maackii (Amur honeysuckle). In tornado sites, honeysuckle thinned out over time, leaving larger shrubs with greater mean basal diameter compared to the undisturbed forest. Other woody invasive species were also more prevalent in the damaged area, but increased in number in both locations by 2017. The forest has the capability to begin to recover from the initial tornado, but its future composition may differ from its initial trajectory due to invasive species, loss of ash trees, and anthropogenic impacts within the urban landscape.

Abandoning conversion from even-aged to uneven-aged forest stands – the effects on production and economic returns

ABSTRACT There is growing interest in continuous cover forestry in Sweden. The historical domination of even-aged forest management means there is a need to focus on methods for transforming even-aged to uneven-aged stands. Practical conversion management today is highly adaptive, and the possibility of failures, such as persistent lack of tree recruitment, must be allowed for. We used simulations to evaluate various scenarios in which conversion management is introduced at different development stages in even-aged stands and then abandoned in favour of clear-felling. A shift towards an inverse J-shaped diameter distribution at the end of the simulations was most evident in northern Sweden and with an early introduction of conversion forestry. The largest losses were incurred in a scenario where the conversion management started with removing larger trees at pre-commercial thinning: it reduced the volume of production by up to 25% and, if clear-felled, resulted in a lower land expectation value than conventional management. Earlier decision to abandon conversion management reduced the losses, but the effect of timing was minor. In summary, the results indicate that conversion management could be started and abandoned without any major economic loss during the timeframe of a normal rotation.

Multi-dimensional temperature sensitivity of protected tropical mountain rain forests

IntroductionTropical mountain rain forests (TMRF, natural forests at > 300 m asl) are globally important for biodiversity and ecosystem services and are believed to be highly vulnerable to climate change. But there are no specific approaches for rigorous assessment of their vulnerability at the landscape and local scales necessary for management for adaptation. We address the challenge of evaluating the ecological sensitivity to temperature of TMRF, applying a multidimensional approach in protected areas over a 440–2,950 m asl altitudinal gradient in Costa Rica, synthesizing results of a long-term research programme (2012-present). We evaluate the sensitivity to the current spatial temperature gradient of eleven ecosystem properties in three categories: forest composition and diversity, thermal characteristics of forest stands and forest structure and dynamics.MethodsData are from 29 to 32 plots of 50 m x 50 m (0.25 ha) distributed over the gradient, in which all trees, palms and tree ferns ≥ 10 dbh are identified to species and measured for recruitment, growth and mortality. An experimental study of leaf litter decomposition rates was carried out in twelve plots. Current and future (SSP 585, 2070) values of mean annual temperatures MAT were obtained from online climate surfaces. Thermal characteristics of forest stands were determined using MATs of species occurrences in GBIF and include a new index, the Community Thermal Capital Index (CTCI), calculated as CTI-MAT.ResultsWe classified degrees of sensitivity to temperature as very weak, weak, moderate or substantial. All eleven ecosystem properties are substantially sensitive, so changes in their values are expected under rising temperatures. Species density, the community temperature index CTI, tree recruitment and mortality rates and leaf litter decomposition rates are positively related to temperature, while the community weighted mean thermal niche breadth, the CTCI, net basal area increments, stand basal area and carbon in aboveground biomass are negatively related. Results point to zones of vulnerability in the protected areas.DiscussionIn montane forests, positive values of the CTCI–climate credit– robust basal area growth and very low mortality and leaf litter decomposition rates suggest healthy ecosystems and no risk of mountaintop extinction. Lowland forests may be vulnerable to degradation and biotic attrition, showing current basal area loss, high mortality and climate debts. National and local actors are participating in a process of adoption of the sensitivity analysis and recommendations regarding zones of vulnerability.

Stand dynamics of old-growth hemlock forests in central Bhutan are shaped by natural disturbances

Understanding how past disturbances have influenced the development of forests is critical for deciphering their current structure and composition and forecasting future changes. In this study, dendrochronological methods were applied to uncover the disturbance history of old-growth hemlock-dominated forests in central Bhutan. Analysis of tree-ring samples from two old-growth hemlock stands, located in two different topographic settings, identified the importance of gap-phase dynamics in facilitating recruitment and growth releases and producing complex, multi-aged structures over time. One site showed evidence of a near stand-replacing disturbance in the late 1700s, while the other showed no evidence of high-severity disturbance at any time over the last 400 years. At both sites low-to medium-severity disturbances, some of which appear to be associated with cyclones originating in the Bay of Bengal, dominated the disturbance regime. The hemlock stands exhibited a significant positive association between cyclone occurrence and growth release events and between recruitment pulses and growth release events. From 1800 to 1970 there was an increase in recruitment of angiosperm tree species at most sites and a corresponding decline in conifer recruitment. Over the past 50 years there has been little new recruitment; this may be due to light limitation in the understory from shade-tolerant angiosperms and bamboo in the lower strata of these stands. Significant variations in disturbance dynamics and recruitment were observed across the study sites, suggesting that other factors, such as topography and climate, may be influencing long-term stand development patterns. This study highlights the complex interplay between historical disturbance regimes and tree recruitment in shaping the age and size structures of old-growth hemlock forests in central Bhutan. It also provides new insights into the dynamics of these forests that can be used to support effective forest conservation and management in the future.

Granivore abundance shapes mutualism quality in plant-scatterhoarder interactions.

Conditional mutualisms involve costs and benefits that vary with environmental factors, but mechanisms driving these dynamics remain poorly understood. Scatterhoarder-plant interactions are a prime example of this phenomenon, as scatterhoarders can either increase or reduce plant recruitment depending on the balance between seed dispersal and predation. We explored factors that drive the magnitude of net benefits for plants in this interaction using a mathematical model, with parameter values based on European beech (Fagus sylvatica) and yellow-necked mice (Apodemus flavicollis). We measured benefits as the percentage of germinating seeds, and examined how varying rodent survival (reflecting, e.g. changes in predation pressure), the rate of seed loss to other granivores, the abundance of alternative food resources, and changes in masting patterns affect the quality of mutualism. We found that increasing granivore abundance can degrade the quality of plant-scatterhoarder mutualism due to increased cache pilferage. Scatterhoarders are predicted to respond by increasing immediate consumption of gathered seeds, leading to higher costs and reduced benefits for plants. Thus, biotic changes that are detrimental to rodent populations can be beneficial for tree recruitment due to adaptive behavior of rodents. When scatterhoarder populations decline too drastically (< 5 individuals ha-1 ); however, tree recruitment may also suffer.

Vegetation dynamics of permanent plot in dipterocarp forest Bukit Bangkirai, East Kalimantan, Indonesia

Vegetation dynamics in dipterocarp forest permanent plots after fire (1997-1998) is important to monitor due to the successional process. Monitoring was conducted from 2001 to 2013 to record tree individuals with a more than 4.8 cm diameter. The highest species diversity was found in 2013 (249 species), then decreased in 2022 (237 species). Dominant species during monitoring change from Macaranga gigantea (Euphorbiaceae) and Madhuca kingiana (Sapotaceae) (2001-2002) to M. gigantea (Euphorbiaceae) and Vernonia arborea (2005-2013) and then M. gigantea (Euphorbiaceae) and M. kingiana (Sapotaceae) (2022). Shannon-Wiener diversity index (H’) in 2022 indicates that species diversity in this permanent plot is high (4.85). The class diameter of trees in the permanent plot is dominated by small individuals (DBH <10 cm). The highest diameter increments are M. gigantea (0.42 cm year-1) and V. arborea (0.41 cm year-1) in the 5-10 cm diameter class. The highest tree recruitment found in 2005 (41.62%). Mortality rate from 2013-2022 was 3.16%, with high contribution from M. gigantea (141 individuals). Carbon storage value from 2001 to 2022 increased from 38.91 tons ha-1 (2001) to 114.29 tons ha-1 (2022). Successional process in this permanent plot needs to be monitored to assess forest health.

Effects of freezing temperatures on early life stages of native trees of different elevational origin: implications for tree recruitment in seasonally dry mountain forests.

In mountain forests, tree regeneration is limited by increasingly frequent frosts with increasing elevation. We investigated the effects of exposure to freezing temperature on early life stages of two native trees of different elevational origin in a seasonally dry mountain forest. We hypothesized that the negative effects of freezing exposure on performance of early life stages increases as freezing temperature decreases, and that frost resistance increases in plants of high elevational origin. We collected seeds of two tree species (Kageneckia lanceolata and Lithraea molleoides) from populations located at different elevations and grew seedlings and saplings in a greenhouse. Dry seeds, imbibed seeds and 1-month-old seedlings were exposed to seven temperature treatments ranging from 4 °C to -20 °C, while 12-month-old saplings were exposed to four temperature treatments from -8 °C to -20 °C. After freezing exposure in a climate chamber, we monitored seed germination and seedling and sapling survival. Germination of K. lanceolata decreased with decreasing temperature only for imbibed seeds from mid- and high elevations, whereas germination of L. molleoides slightly increased with decreasing temperature only for imbibed seeds from high elevations. For both species, seedling survival decreased with decreasing temperature. For K. lanceolata, the negative effects of freezing temperatures were weaker as elevational origin of seeds increased, whereas L. molleoides showed the opposite pattern. For both species, saplings only survived at the mildest applied freezing temperature (-8 °C). We conclude that effects of climatic variation associated with elevation depend on the study species and life stage. The observed patterns could be caused by maternal effects, which are absent at the sapling stage. Moreover, temperatures below -8 °C can limit recruitment since partial mortality of seedlings and saplings occurred at such values.

Regeneration dynamics in mixed mountain forests at their natural geographical distribution range in the Western Rhodopes

Mixed mountain forests consisting of Norway spruce (Picea abies (L.) Karst.), European beech (Fagus sylvatica L.), and silver fir (Abies alba Mill.) are among the most productive and stable forest ecosystems in Europe. Their southeasternmost geographical distribution range is located in the Western Rhodopes, where they have high economic, recreational, and ecological value. In the past, shelterwood cuttings dominated forest management practices in these forests and were mainly aimed at maintaining and reproducing conifers. During the past two decades, single-tree and group-tree selection systems have been promoted as alternative management approaches to support the conversion of spruce-dominated stands to close-to-nature mixed forests of fir, beech, and spruce. However, the natural regeneration dynamics in these stands are barely known, and their dependence on microsite and management effects needs to be better understood.The objective of this study was to investigate ecological factors under management regimes of different intensity (“single-tree”-selection and “group-tree”-selection) that influence the regeneration processes in mixed mountain forests in the Bulgarian Rhodopes. Data on regeneration and microsite conditions were collected on 105 systematically distributed plots (25 m2/100 m2) in four 100–120 years old stands located in the regional forest district of Smoljan, Bulgaria (1580–1650 m a.s.l.). We relied on generalizeds linear mixed models to analyse for each species the (1) size-dependent regeneration density and (2) height increment in dependence on management practices, competing vegetation, as well as soil and light conditions.Our study revealed an overall high potential for recruitment in the Western Rhodopes. Regeneration density was highest in fir (median 12800 N ha-1), followed by spruce (median 1600 N ha-1) and beech (median 1200 N ha-1). Fir benefited most from “single-tree” selection cuttings, while “group-tree” selection cutting tended to promote beech and fir but also spruce. Competing ground vegetation was detrimental for seedling density of all species. Annual height increment increased with plant size, was lowest in spruce, and similar in fir and beech. Sapling increment was driven by light, whereas seedlings did not react to increased radiation. Browsing was species-specific and was highest in beech (15–30 %), followed by fir (5–10 %) and spruce (<1 %). It was not a crucial factor in impeding tree recruitment. We conclude that frequent harvest activities of low intensity which consider advanced regeneration are a promising approach to successfully convert the formerly spruce-fir-dominated forests to climate-adapted fir-beech-(spruce)-mixed stands.

Forest recovery through applied nucleation: Effects of tree islet size and disperser mobility on tree recruitment in a temperate landscape

The plantation of tree islets within deforested areas (applied nucleation) has been proposed as a restoration strategy able to catalyze tree colonization in the matrix with reasonable costs and labor compared to traditional restoration methods such as extensive plantations. However, recent work has questioned its effectiveness in comparison to natural forest regeneration, and observational studies have eventually reported a limited expansion of tree islets after years of planting. To infer the processes affecting the effectiveness of applied nucleation, we used a stochastic model able to predict avian movement and fruit consumption, tree seed dispersal and seedling establishment in a fragmented temperate forest landscape of northern Spain. First, we simulated scenarios of natural regeneration and compared them to applied nucleation in terms of tree colonization in the remaining deforested matrix. Subsequently, in applied nucleation scenarios we assessed the effects of islet size and matrix area recovered on tree recruitment into the matrix. Also, we evaluated if such effects were mediated by the scale of movement of frugivores. Our results showed that applied nucleation triggers matrix colonization by trees at higher rates than natural regeneration because it avoids frugivore entrapment within forest edges. Also, that large tree islets can hamper matrix colonization despite being visited more frequently. If large tree islets offer abundant resources, animals do not need to leave their boundaries, concentrating seed deposition within islets. As expected, such effects depended on the scale of movement of frugivores. Our work highlights that in nucleated landscapes, islets harboring fruit-producing trees should attract animal dispersers, and also foster animal inter-islet movement. Both conditions should be met to ensure seed deposition and seedling establishment in the deforested matrix, promoting forest recovery independent of islet expansion through vegetative growth. To achieve these restoration goals, we propose establishing islets whose size and inter-islet distances matches the scale of movement of main seed dispersers present in the area.