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With ongoing global warming, increasing water deficits promote physiological stress on forest ecosystems with negative impacts on tree growth, vitality, and survival. How individual tree species will react to increased drought stress is therefore a key research question to address for carbon accounting and the development of climate change mitigation strategies. Recent tree-ring studies have shown that trees at higher latitudes will benefit from warmer temperatures, yet this is likely highly species-dependent and less well-known for more temperate tree species. Using a unique pan-European tree-ring network of 26,430 European beech (Fagus sylvatica L.) trees from 2118 sites, we applied a linear mixed-effects modeling framework to (i) explain variation in climate-dependent growth and (ii) project growth for the near future (2021-2050) across the entire distribution of beech. We modeled the spatial pattern of radial growth responses to annually varying climate as a function of mean climate conditions (mean annual temperature, mean annual climatic water balance, and continentality). Over the calibration period (1952-2011), the model yielded high regional explanatory power (R 2  = 0.38-0.72). Considering a moderate climate change scenario (CMIP6 SSP2-4.5), beech growth is projected to decrease in the future across most of its distribution range. In particular, projected growth decreases by 12%-18% (interquartile range) in northwestern Central Europe and by 11%-21% in the Mediterranean region. In contrast, climate-driven growth increases are limited to around 13% of the current occurrence, where the historical mean annual temperature was below ~6°C. More specifically, the model predicts a 3%-24% growth increase in the high-elevation clusters of the Alps and Carpathian Arc. Notably, we find little potential for future growth increases (-10 to +2%) at the poleward leading edge in southern Scandinavia. Because in this region beech growth is found to be primarily water-limited, a northward shift in its distributional range will be constrained by water availability., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

The fundamental trade-off between current and future reproduction has long been considered to result in a tendency for species that can grow large to begin reproduction at a larger size. Due to the prolonged time required to reach maturity, estimates of tree maturation size remain very rare and we lack a global view on the generality and the shape of this trade-off. Using seed production from five continents, we estimate tree maturation sizes for 486 tree species spanning tropical to boreal climates. Results show that a species' maturation size increases with maximum size, but in a non-proportional way: the largest species begin reproduction at smaller sizes than would be expected if maturation were simply proportional to maximum size. Furthermore, the decrease in relative maturation size is steepest in cold climates. These findings on maturation size drivers are key to accurately represent forests' responses to disturbance and climate change., (© 2024 John Wiley & Sons Ltd.)

Natural forest expansion (NFE), that is, the establishment of secondary forest on non-forested land through natural succession, has substantially contributed to the widespread expansion of forests in Europe over the last few decades. So far, EU policies have largely neglected the potential of NFE for meeting policy objectives on restoration. Synthesising recent interdisciplinary research, this paper assesses the challenges and opportunities of NFE in view of contributing to European forest and ecosystem restoration. Specifically, we discuss the potential for supporting climate change mitigation and adaptation, biodiversity conservation, and forestry and economic use, summarize the current knowledge about societal perceptions and the policymaking on NFE, and make policy recommendations to better use the potential of NFE. We conclude that NFE has the potential to contribute to the European restoration policy agenda if local contexts and possible trade-offs are properly considered., (© 2023. The Author(s).)

The periodic production of large seed crops, or masting, is a widespread phenomenon in perennial plants. This behavior can enhance the reproductive efficiency of plants, leading to increased fitness, and produce ripple effects on food webs. While variability from year to year is a defining characteristic of masting, the methods used to quantify this variability are highly debated. The commonly used coefficient of variation lacks the ability to account for the serial dependence in mast data and can be influenced by zeros, making it a less suitable choice for various applications based on individual-level observations, such as phenotypic selection, heritability, and climate change studies, which rely on individual-plant-level datasets that often contain numerous zeros. To address these limitations, we present three case studies and introduce volatility and periodicity, which account for the variance in the frequency domain by emphasizing the significance of long intervals in masting. By utilizing examples of Sorbus aucuparia, Pinus pinea, Quercus robur, Quercus pubescens, and Fagus sylvatica, we demonstrate how volatility captures the effects of variance at both high and low frequencies, even in the presence of zeros, leading to improved ecological interpretations of the results. The growing availability of long-term, individual-plant datasets promises significant advancements in the field, but requires appropriate tools for analysis, which the new metrics provide., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

The tropical Andes are one of the most important biodiversity hotspots on Earth, yet our understanding of how their biotas have responded to Quaternary climatic oscillations is extraordinarily limited and the alternative models proposed to explain their demographic dynamics have been seldom formally evaluated. Here, we test the hypothesis that the interplay between the spatial configuration of geographical barriers to dispersal and elevational displacements driven by Quaternary cooling-warming cycles has shaped the demographic trajectories of montane oak forests (Quercus humboldtii) from the Colombian Andes. Specifically, we integrate genomic data and environmental niche modelling at fine temporal resolution to test competing spatially explicit demographic and coalescent models, including scenarios considering (i) isotropic gene flow through the landscape, (ii) the hypothetical impact of contemporary barriers to dispersal (i.e., inter-Andean valleys), and (iii) distributional shifts of montane oak forests from the Last Glacial Maximum to the present. Although our data revealed a marked genetic fragmentation of montane oak forests, statistical support for isolation-with-migration models indicates that geographically separated populations from the different Andean Cordilleras regularly exchange gene flow. Accordingly, spatiotemporally explicit demographic analyses supported a model of flickering connectivity, with scenarios considering isotropic gene flow or currently unsuitable habitats as persistent barriers to dispersal providing a comparatively worse fit to empirical genomic data. Overall, these results emphasize the role of landscape heterogeneity on shaping spatial patterns of genomic variation in montane oak forests, rejecting the hypothesis of genetic continuity and supporting a significant impact of Quaternary climatic oscillations on their demographic trajectories., (© 2023 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

Background and Aims: In a range of plant species, the distribution of individual mean fecundity is skewed and dominated by a few highly fecund individuals. Larger plants produce greater seed crops, but the exact nature of the relationship between size and reproductive patterns is poorly understood. This is especially clear in plants that reproduce by exhibiting synchronized quasi-periodic variation in fruit production, a process called masting., Methods: We investigated covariation of plant size and fecundity with individual-plant-level masting patterns and seed predation in 12 mast-seeding species: Pinus pinea, Astragalus scaphoides, Sorbus aucuparia, Quercus ilex, Q. humilis, Q. rubra, Q. alba, Q. montana, Chionochloa pallens, C. macra, Celmisia lyallii and Phormium tenax., Key Results: Fecundity was non-linearly related to masting patterns. Small and unproductive plants frequently failed to produce any seeds, which elevated their annual variation and decreased synchrony. Above a low fecundity threshold, plants had similar variability and synchrony, regardless of their size and productivity., Conclusions: Our study shows that within-species variation in masting patterns is correlated with variation in fecundity, which in turn is related to plant size. Low synchrony of low-fertility plants shows that the failure years were idiosyncratic to each small plant, which in turn implies that the small plants fail to reproduce because of plant-specific factors (e.g. internal resource limits). Thus, the behaviour of these sub-producers is apparently the result of trade-offs in resource allocation and environmental limits with which the small plants cannot cope. Plant size and especially fecundity and propensity for mast failure years play a major role in determining the variability and synchrony of reproduction in plants., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Recruitment is a primary determinant of the long-term dynamics of plant populations in changing environments. However, little information is known about the effects of anthropogenic environmental changes on reproductive ecology of trees. We evaluated the impact of experimentally induced 18 yr of drought on reproduction of three contrasting forest trees: Quercus ilex, Phillyrea latifolia and Arbutus unedo. Rainfall reduction did not decrease tree fecundity. Drought, however, affected the allocation of resources in Q. ilex and A. unedo but not the more drought tolerant P. latifolia. Larger crop production by Q. ilex and A. unedo was associated with a stronger decrease in growth in the rainfall-reduction plots compared with the control plots, suggesting that these species were able to maintain their fecundity by shifting their allocation of resources away from growth. Our results indicated resistance to change in tree fecundity in Mediterranean-type forest subjected to an average 15% decrease in the amount of soil moisture, suggesting that these ecosystems may adapt to a progressive increase in arid conditions. However, the species-specific reductions in growth may indirectly affect future fecundity and ultimately shift community composition, even without immediate direct effects of drought on tree fecundity., (© 2020 The Authors. New Phytologist © 2020 New Phytologist Trust.)

Masting-temporally variable seed production with high spatial synchrony-is a pervasive strategy in wind-pollinated trees that is hypothesized to be vulnerable to climate change due to its correlation with variability in abiotic conditions. Recent work suggests that aging may also have strong effects on seed production patterns of trees, but this potential confounding factor has not been considered in previous times series analysis of climate change effects. Using a 54 year dataset for seven dominant species in 17 forests across Poland, we used the proportion of seed-producing trees (PST) to contrast the predictions of the climate change and aging hypotheses in Abies alba, Fagus sylvatica, Larix decidua, Picea abies, Pinus sylvestris, Quercus petraea, and Quercus robur. Our results show that in all species, PST increased over time and that this change correlated most strongly with stand age, while the standardized precipitation-evapotranspiration index, a measure of drought, contributed to temporal trends in PST of F. sylvatica and Q. robur. Temporal variability of PST also increased over time in all species except P. sylvestris, while trends in temporal autocorrelation and among-stand synchrony reflect species-specific masting strategies. Our results suggest a pivotal role of plant ontogeny in driving not only the extent but also variability and synchrony of reproduction in temperate forest trees. In a time of increasing forest regrowth in Europe, we therefore call for increased attention to demographic effects such as aging on plant reproductive behavior, particularly in studies examining global change effects using long-term time series data., (© 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Mast seeding is one of the most intriguing reproductive traits in nature. Despite its potential drawbacks in terms of fitness, the widespread existence of this phenomenon suggests that it should have evolutionary advantages under certain circumstances. Using a global dataset of seed production time series for 219 plant species from all of the continents, we tested whether masting behaviour appears predominantly in species with low foliar nitrogen and phosphorus concentrations when controlling for local climate and productivity. Here, we show that masting intensity is higher in species with low foliar N and P concentrations, and especially in those with imbalanced N/P ratios, and that the evolutionary history of masting behaviour has been linked to that of nutrient economy. Our results support the hypothesis that masting is stronger in species growing under limiting conditions and suggest that this reproductive behaviour might have evolved as an adaptation to nutrient limitations and imbalances.

Synchronized and variable reproduction by perennial plants, called mast seeding, is a major reproductive strategy of trees. The need to accumulate sufficient resources after depletion following fruiting (resource budget), the efficiency of mass flowering for outcross pollination (pollen coupling), or the external factors preventing reproduction (environmental veto) could all synchronize masting. We used seed production data for four species ( Quercus ilex , Quercus humilis , Sorbus aucuparia , and Pinus albicaulis ) to parametrize resource budget models of masting. Based on species life-history characteristics, we hypothesized that pollen coupling should synchronize reproduction in S. aucuparia and P. albicaulis , while in Q. ilex and Q. humilis , environmental veto should be a major factor. Pollen coupling was stronger in S. aucuparia and P. albicaulis than in oaks, while veto was more frequent in the latter. Yet in all species, costs of reproduction were too small to impose a replenishment period. A synchronous environmental veto, in the presence of environmental stochasticity, was sufficient to produce observed variability and synchrony in reproduction. In the past, vetoes like frost events that prevent reproduction have been perceived as negative for plants. In fact, they could be selectively favored as a way to create mast seeding.

Background and Aims: The ability of plants to allocate energy to resistance against herbivores changes with abiotic conditions and thus may vary along geographical clines, with important consequences for plant communities. Seed size is a plant trait potentially influencing plant tolerance to endoparasites, and seed size often varies across latitude. Consequently, plant tolerance to endoparasites may change across geographical clines., Methods: The interaction between Quercus ilex (holm oak) and seed-predating Curculio spp. (weevils) was explored along most of the latitudinal range of Q. ilex. This included quantification of variation in seed size, survival likelihood of infested seeds, multi-infestation of acorns and community composition of Curculio weevils in acorns., Key Results: Larger seeds had a higher probability of surviving weevil attack (i.e. embryo not predated). Southern populations of oak produced on average four times larger seeds than those of northern populations. Consequently, the probability of survival of infested acorns decreased with latitude. The community composition of Curculio varied, with large weevils (C. elephas) dominating in southern populations and small weevils (C. glandium) dominating in northern populations. However, damage tolerance was robust against this turnover in predator functional traits. Furthermore, we did not detect any change in multi-infestation of acorns along the geographical gradient., Conclusions: Quercus ilex tolerance to seed predation by Curculio weevils increases toward the southern end of its distribution. Generally, studies on geographical variation in plant defence against enemies largely ignore seed attributes or they focus on seed physical barriers. Thus, this research suggests another dimension in which geographical trends in plant defences should be considered, i.e. geographical variation in tolerance to seed predators mediated by seed size., (© The Author(s) 2018. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Species introduced to habitats outside their native range often escape control by their natural enemies. Besides competing with native species, an alien species might also affect the native herbivores by introducing a new source of different quality food. Here, we describe the case of northern red oak (Quercus rubra) invasion in Europe. We collected data on insect (moth Cydia spp. and weevil Curculio spp.) seed predation of northern red oak in its native (USA, North America) and invasive (Poland, Europe) range, as well as for sessile oaks (Quercus petrea) in Europe. We also evaluated the quality of acorns as hosts for weevil larvae by collecting infested acorns and measuring weevil developmental success, and quantifying acorn traits such as seed mass, tannins, lipids and protein concentration. We used DNA barcoding to identify insects to the species level. The predation by moths was similar and very low in both species and in both ranges. However, red oaks escape pre-dispersal seed predation by weevils in Europe. Weevil infestation rates of northern red oak acorns in their invasive range were 10 times lower than that of sessile oaks, and also 10 times lower than that of red oaks in North America. Furthermore, even when weevils oviposited into northern red oaks, the larvae failed to develop, suggesting that the exotic host created a trap for the insect. This phenomenon might gradually decrease the local abundance of the seed predator, and further aid the invasion., (© 2017 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

The function and conservation of many forest ecosystems depend on the distribution and diversity of the community of rodents that consume and disperse seeds. The habitat preferences and interactions are especially relevant in alpine systems where such granivorous rodents reach the southernmost limit of their distribution and are especially sensitive to global warming. We analyzed the community of granivorous rodents in the Pyrenees, one of the southernmost mountain ranges of Europe. Rodent species were identified by DNA with particular attention to the Apodemus species, which are prominent seed-dispersing rodents in Europe. We confirmed for the first time the presence of the yellow-necked mouse, Apodemus flavicollis, in central Pyrenees, a typical Eurosiberian species that reaches its southernmost distribution limit in this area. We also found the wood mouse, Apodemus sylvaticus, a related species more tolerant to Mediterranean environments. Both rodents were spatially segregated by altitude. A. sylvaticus was rare at high altitudes, which might cause the genetic differentiation between populations of the different valleys reported here. We also found other seed consumers like dormice, Elyomis quercinus, and voles, Myodes glareolus, with marked habitat preferences. We suggest that population isolation among valleys may increase the genetic diversity of rodents, like A. sylvaticus. We also highlight the potential threat that global warming may represent for species linked to high-altitude refuges at the southern edge of its distribution, like Apodemus flavicollis. Finally, we discuss how this threat may have a dimension in the conservation of alpine forests dispersed by these rodent populations., (© 2018 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Synchrony between seed growth and oogenesis is suggested to largely shape trophic breadth of seed-feeding insects and ultimately to contribute to their co-existence by means of resource partitioning or in the time when infestation occurs. Here we investigated: (i) the role of seed phenology and sexual maturation of females in the host specificity of seed-feeding weevils (Curculio spp.) predating in hazel and oak mixed forests; and (ii) the consequences that trophic breadth and host distribution have in the genetic structure of the weevil populations. DNA analyses were used to establish unequivocally host specificity and to determine the population genetic structure. We identified 4 species with different specificity, namely Curculio nucum females matured earlier and infested a unique host (hazelnuts, Corylus avellana) while 3 species (Curculio venosus, Curculio glandium and Curculio elephas) predated upon the acorns of the 2 oaks (Quercus ilex and Quercus pubescens). The high specificity of C. nucum coupled with a more discontinuous distribution of hazel trees resulted in a significant genetic structure among sites. In addition, the presence of an excess of local rare haplotypes indicated that C. nucum populations went through genetic expansion after recent bottlenecks. Conversely, these effects were not observed in the more generalist Curculio glandium predating upon oaks. Ultimately, co-existence of weevil species in this multi-host-parasite system is influenced by both resource and time partitioning. To what extent the restriction in gene flow among C. nucum populations may have negative consequences for their persistence in a time of increasing disturbances (e.g. drought in Mediterranean areas) deserves further research., (© 2017 The Authors. Integrative Zoology published by International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Variation in seed availability shapes plant communities, and is strongly affected by seed predation. In some plant species, temporal variation in seed production is especially high and synchronized over large areas, which is called 'mast seeding'. One selective advantage of this phenomenon is predator satiation which posits that masting helps plants escape seed predation through starvation of predators in lean years, and satiation in mast years. However, even though seed predation can be predicted to have a strong spatial component and depend on plant densities, whether the effectiveness of predator satiation in masting plants changes according to the Janzen-Connell effect has been barely investigated. We studied, over an 8-year period, the seed production, the spatiotemporal patters of weevil seed predation, and the abundance of adult weevils in a holm oak (Quercus ilex) population that consists of trees interspersed at patches covering a continuum of conspecific density. Isolated oaks effectively satiate predators, but this is trumped by increasing conspecific plant density. Lack of predator satiation in trees growing in dense patches was caused by re-distribution of insects among plants that likely attenuated them against food shortage in lean years, and changed the type of weevil functional response from type II in isolated trees to type III in trees growing in dense patches. This study provides the first empirical evaluation of the notion that masting and predator satiation should be more important in populations that start to dominate their communities, and is consistent with the observation that masting is less frequent and less intense in diverse forests.

Masting is the highly variable production of synchronized seed crops, and is a common reproductive strategy in plants. Weather has long been recognized as centrally involved in driving seed production in masting plants. However, the theory behind mechanisms connecting weather and seeding variation has only recently been developed, and still lacks empirical evaluation. We used 12-year long seed production data for 255 holm oaks ( Quercus ilex ), as well as airborne pollen and meteorological data, and tested whether masting is driven by environmental constraints: phenological synchrony and associated pollination efficiency, and drought-related acorn abscission. We found that warm springs resulted in short pollen seasons, and length of the pollen seasons was negatively related to acorn production, supporting the phenological synchrony hypothesis. Furthermore, the relationship between phenological synchrony and acorn production was modulated by spring drought, and effects of environmental vetoes on seed production were dependent on last year's environmental constraint, implying passive resource storage. Both vetoes affected among-tree synchrony in seed production. Finally, precipitation preceding acorn maturation was positively related to seed production, mitigating apparent resource depletion following high crop production in the previous year. These results provide new insights into mechanisms beyond widely reported weather and seed production correlations., (© 2017 The Author(s).)

Host specialization has long been hypothesized to explain the extraordinary diversity of phytophagous insects in the tropics. However, addressing this hypothesis has proved challenging because of the risk of over-looking rare interactions, and hence biasing specialization estimations, and the difficulties to separate the diversity component attributable to insect specialization from that related to host diversity. As a result, the host specialization hypothesis lacks empirical support for important phytophagous insect clades. Here, we test the hypothesis in a radiation of seed-feeding insects, acorn weevils (Curculio spp.), sampled in temperate and tropical regions (California and Nicaragua, respectively) with an equivalent pool of oak host species. Using DNA sequences from three low-copy genes, we delimited to species level 778 weevil larvae extracted from host seeds and assessed their phylogenetic relationships by Maximum Likelihood and Bayesian inference. We then reconstructed the oak-weevil food webs and examined differences in alpha, beta and gamma diversity using Hill numbers of effective species. We found a higher alpha, beta and gamma diversity of weevils in Nicaragua compared to California despite similar richness of host species at both local and regional level. By means of Bayesian mixed models, we also found that tropical weevil species were highly specialized both in terms of host range and interaction strength, whereas their temperate congeners had a broader taxonomic and phylogenetic host spectrum. Finally, in Nicaraguan species, larval body size was highly correlated with the size of the acorns infested, as would be expected by a greater host specialization, whereas in California this relationship was absent. Altogether, these lines of evidence support the host specialization hypothesis and suggest contrasting eco-evolutionary dynamics in tropical and temperate regions even in absence of differences in host diversity., (© 2017 by the Ecological Society of America.)

Fruit production (NPP f ), the amount of photosynthates allocated to reproduction (%GPP f ) and their controls for spatial and species-specific variability (e.g. nutrient availability, climate) have been poorly studied in forest ecosystems. We characterized fruit production and its temporal behaviour for several tree species and resolved the effects of gross primary production (GPP), climate and foliar nutrient concentrations. We used data for litterfall and foliar nutrient concentration from 126 European forests and related them to climatic data. GPP was estimated for each forest using a regression model. Mean NPP f ranged from c. 10 to 40 g C m -2  yr -1 and accounted for 0.5-3% of GPP. Forests with higher GPPs produced larger fruit crops. Foliar zinc (Zn) and phosphorus (P) concentrations were associated positively with NPP f , whereas foliar Zn and potassium (K) were negatively related to its temporal variability. Maximum NPP f and interannual variability of NPP f were higher in Fagaceae than in Pinaceae species. NPP f and %GPP f were similar amongst the studied species despite the different reproductive temporal behaviour of Fagaceae and Pinaceae species. We report that foliar concentrations of P and Zn are associated with %GPP f , NPP f and its temporal behaviour., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

The complexity of animal life histories makes it difficult to predict the consequences of climate change on their populations. In this paper, we show, for the first time, that longer summer drought episodes, such as those predicted for the dry Mediterranean region under climate change, may bias insect population sex ratio. Many Mediterranean organisms, like the weevil Curculio elephas, become active again after summer drought. This insect depends on late summer rainfall to soften the soil and allow adult emergence from their underground refuges. We found that, as in many protandric species, more C. elephas females emerged later in the season. Male emergence timing was on average earlier and also more dependent on the beginning of late summer rainfall. When these rains were delayed, the observed weevil sex ratio was biased towards females. So far, the effects of global warming on animal sex ratios has been reported for temperature-dependent sex determination in reptiles. Our results show that rainfall timing can also bias the sex ratio in an insect, and highlight the need for keeping a phenological perspective to predict the consequences of climate change. We must consider not just the magnitude of the predicted changes in temperature and rainfall but also the effects of their timing.

The patterns of seedling recruitment in animal-dispersed plants result from the interactions among environmental and behavioral variables. However, we know little on the contribution and combined effect of both kinds of variables. We designed a field study to assess the interplay between environment (vegetation structure, seed abundance, rodent abundance) and behavior (seed dispersal and predation by rodents, and rooting by wild boars), and their contribution to the spatial patterns of seedling recruitment in a Mediterranean mixed-oak forest. In a spatially explicit design, we monitored intensively all environmental and behavioral variables in fixed points at a small spatial scale from autumn to spring, as well as seedling emergence and survival. Our results revealed that the spatial patterns of seedling emergence were strongly related to acorn availability on the ground, but not by a facilitation effect of vegetation cover. Rodents changed seed shadows generated by mother trees by dispersing most seeds from shrubby to open areas, but the spatial patterns of acorn dispersal/predation had no direct effect on recruitment. By contrast, rodents had a strong impact on recruitment as pilferers of cached seeds. Rooting by wild boars also reduced recruitment by reducing seed abundance, but also by changing rodent's behavior towards higher consumption of acorns in situ. Hence, seed abundance and the foraging behavior of scatter-hoarding rodents and wild boars are driving the spatial patterns of seedling recruitment in this mature oak forest, rather than vegetation features. The contribution of vegetation to seedling recruitment (e.g. facilitation by shrubs) may be context dependent, having a little role in closed forests, or being overridden by directed seed dispersal from shrubby to open areas. We warn about the need of using broad approaches that consider the combined action of environment and behavior to improve our knowledge on the dynamics of natural regeneration in forests.

Understanding the factors determining genetic diversity and structure in peripheral populations is a long-standing goal of evolutionary biogeography, yet little empirical information is available for tropical species. In this study, we combine information from nuclear microsatellite markers and niche modelling to analyse the factors structuring genetic variation across the southernmost populations of the tropical oak Quercus segoviensis. First, we tested the hypothesis that genetic variability decreases with population isolation and increases with local habitat suitability and stability since the Last Glacial Maximum (LGM). Second, we employed a recently developed multiple matrix regression with randomisation (MMRR) approach to study the factors associated with genetic divergence among the studied populations and test the relative contribution of environmental and geographic isolation to contemporary patterns of genetic differentiation. We found that genetic diversity was negatively correlated with average genetic differentiation with other populations, indicating that isolation and limited gene flow have contributed to erode genetic variability in some populations. Considering the relatively small size of the study area (<120 km), analyses of genetic structure indicate a remarkable inter-population genetic differentiation. Environmental dissimilarity and differences in current and past climate niche suitability and their additive effects were not associated with genetic differentiation after controlling for geographic distance, indicating that local climate does not contribute to explain spatial patterns of genetic structure. Overall, our data indicate that geographic isolation, but not current or past climate, is the main factor determining contemporary patterns of genetic diversity and structure within the southernmost peripheral populations of this tropical oak., (© 2014 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Predicting climate-driven changes in plant distribution is crucial for biodiversity conservation and management under recent climate change. Climate warming is expected to induce movement of species upslope and towards higher latitudes. However, the mechanisms and physiological processes behind the altitudinal and latitudinal distribution range of a tree species are complex and depend on each tree species features and vary over ontogenetic stages. We investigated the altitudinal distribution differences between juvenile and adult individuals of seven major European tree species along elevational transects covering a wide latitudinal range from southern Spain (37°N) to northern Sweden (67°N). By comparing juvenile and adult distributions (shifts on the optimum position and the range limits) we assessed the response of species to present climate conditions in relation to previous conditions that prevailed when adults were established. Mean temperature increased by 0.86 °C on average at our sites during the last decade compared with previous 30-year period. Only one of the species studied, Abies alba, matched the expected predictions under the observed warming, with a maximum abundance of juveniles at higher altitudes than adults. Three species, Fagus sylvatica, Picea abies and Pinus sylvestris, showed an opposite pattern while for other three species, such as Quercus ilex, Acer pseudoplatanus and Q. petraea, we were no able to detect changes in distribution. These findings are in contrast with theoretical predictions and show that tree responses to climate change are complex and are obscured not only by other environmental factors but also by internal processes related to ontogeny and demography., (© 2013 John Wiley & Sons Ltd.)

Trophic interactions can trigger the development of exaggerated specialized characters and promote morphological diversification. For example, acorn weevils (genus Curculio) present strikingly long rostrums, which are used by females to perforate oviposition holes through the seed coat. Species exhibiting longer rostrums are known to exploit larger acorns, and therefore rostrum length is thought to be subject to selection to match the preferred acorn type. However, rostrum length is strongly correlated with body size, and morphological divergence could result from either selection on rostrum length for optimal food exploitation or from other pressures acting on body size. We collected infested acorns at oak forests where the large Curculio elephas and the small-bodied Curculio glandium co-occur. There were no interspecific differences in adult female body size to rostrum length allometric relationships, and rostrum length is equally correlated with body size in either species. MtDNA-based species identification showed that C. glandium larvae were present within acorns of all sizes, whereas C. elephas larvae were restricted to acorns above a minimum size, irrespective of oak species. Hence, exploitation of large acorns can hardly have triggered rostrum enlargement, as the small sized C. glandium adults (with short rostrums) could perforate and oviposit in both small and large acorns. Rather, increased rostrum length is probably a by-product of the larger body sizes of individuals emerging from bigger acorns, which allow increased larval size and enhance larval survival likelihood. Summarizing, when exaggerated feeding traits co-vary with other body features, interspecific morphological variability may result from contrasting selective pressures acting on these correlated characters.

The concept of trade-offs between reproduction and other fitness traits is a fundamental principle of life history theory. For many plant species, the cost of sexual reproduction affects vegetative growth in years of high seed production through the allocation of resources to reproduction at different hierarchical levels of canopy organization. We have examined these tradeoffs at the shoot and branch level in an endemic California oak, Quercus lobata, during a mast year. To determine whether acorn production caused a reduction in vegetative growth, we studied trees that were high and low acorn producers, respectively. We observed that in both low and high acorn producers, shoots without acorns located adjacent to reproductive shoots showed reduced vegetative growth but that reduced branch-level growth on acorn-bearing branches occurred only in low acorn producers. The availability of local resources, measured as previous year growth, was the main factor determining acorn biomass. These findings show that the costs of reproduction varied among hierarchical levels, suggesting some degree of physiological autonomy of shoots in terms of acorn production. Costs also differed among trees with different acorn crops, suggesting that trees with large acorn crops had more available resources to allocate for growth and acorn production and to compensate for immediate local costs of seed production. These findings provide new insight into the proximate mechanisms for mast-seeding as a reproductive strategy.

The contribution of pre-dispersal seed predation to inter-specific differences in recruitment remains elusive. In species with no resistance mechanisms, differences in pre-dispersal predation may arise from differences in seed abundance (plant satiation) or in the ability of seeds to survive insect infestation (seed satiation). This study aimed to analyse the impact of pre-dispersal acorn predation by weevils in two co-occurring Mediterranean oaks (Quercus ilex and Quercus humilis) and to compare its relevance with other processes involved in recruitment. We monitored the patterns of acorn production and acorn infestation by weevils and we conducted experimental tests of acorn germination after weevil infestation, post-dispersal predation and seedling establishment in mixed forests. Monitoring and experimental data were integrated in a simulation model to test for the effects of pre-dispersal predation in recruitment. In both oaks pre-dispersal acorn infestation decreased with increasing acorn crop size (plant satiation). This benefited Q. ilex which exhibited stronger masting behaviour than Q. humilis, with almost a single and outstanding reproductive event in 6 years. Acorn infestation was more than twice as high in Q. humilis (47.0%) as in Q. ilex (20.0%) irrespective of the number of seeds produced by each species. Although germination of infested acorns (seed satiation) was higher in Q. humilis (60%) than in Q. ilex (21%), this could barely mitigate the higher infestation rate in the former species, to reduce seed loss. Conversely to pre-dispersal predation, no inter-specific differences were observed either in post-dispersal predation or seedling establishment. Our results indicate that pre-dispersal predation may contribute to differences in seed supply, and ultimately in recruitment, between co-existing oaks. Moreover, they suggest that seed satiation can barely offset differences in seed infestation rates. This serves as a warning against overemphasising seed satiation as a mechanism to overcome seed predation by insects.

Temporally variable production of seed crops by perennial plants (masting) has been hypothesized to be a valuable mechanism in the reduction of seed predation by satiating and starving seed consumers. To achieve these benefits, coexisting species subjected to the same predator would benefit from a similar pattern of seeding fluctuation over time that could lead to a reduction in predation at the within-species level. We tested for the existence of an environmental factor enforcing synchrony in acorn production in two sympatric Mediterranean oaks (Quercus ilex and Q. humilis) and the consequences on within-species and between-species acorn predation, by monitoring 15 mixed forests (450 trees) over seven years. Acorn production in Q. ilex and Q. humilis was highly variable among years, with high population variability (CVp) values. The two species exhibited a very different pattern across years in their initial acorn crop size (sum of aborted, depredated, and sound acorns). Nevertheless, interannual differences in summer water stress modified the likelihood of abortion during acorn ripening and enforced within- and, particularly, between-species synchrony and population variability in acorn production. The increase in CVp from initial to mature acorn crop (after summer) accounted for 33% in Q. ilex, 59% in Q. humilis, and 60% in the two species together. Mean yearly acorn pre-dispersal predation by invertebrates was considerably higher in Q. humilis than in Q. ilex. Satiation and starvation of predators was recorded for the two oaks, and this effect was increased by the year-to-year variability in the size of the acorn crop of the two species combined. Moreover, at a longer time scale (over seven years), we observed a significant reduction in the mean proportion of acorns depredated for each oak and the variability in both species' acorn production combined. Therefore, our results demonstrate that similar patterns of seeding fluctuation over time in coexisting species mediated by an environmental cue (summer drought) may contribute to the reduction of the impact of seed predation at a within-species level. Future research should be aimed at addressing whether this process could be a factor assisting in the coexistence of Q. ilex and Q. humilis.