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Anthropic activities have modelled and simplified southern European forest landscapes for centuries. Over recent decades, new drivers related to human-mediated global change have induced the redistribution of tree species and an increase in more complex forests. However, the current large-scale patterns and drivers of these changes are yet to be fully described for the Mediterranean Basin. In this frame, this work identifies and examines changes in dominance and composition from pure to mixed forests across bioclimatic gradients and forest types in Iberian forests over recent decades based on data from the Spanish National Forest Inventory from 1960 to 2020. Then, considering different environmental, anthropic, and disturbance variables we also identify some of the most important drivers associated with the shifts observed from 1986 to 2020. Our results confirm an ongoing increase in mixed forests involving the replacement of conifers by broadleaved species. These shifts are greater in the Atlantic biogeoregion and in pure broadleaved deciduous forests. Climate warming-associated disturbances such as drought severity together with land use legacies and forest types showed the strongest relationships with the observed changes in the studied forests. Our results support the premise put forward by palaeoecologists which states that the increase in tree mixtures is a natural process reversing the historical human-induced simplification of Iberian forests. The increasing importance of mixed forest in southern Europe makes decisive the revision of forest classifications as well as forest management and conservation plans in order to include these increasingly abundant novel stands in forest policies.

The reproductive success of plants depends both on their phenotype and the local neighbourhood in which they grow. Animal-pollinated plants may benefit from increased visitation when surrounded by attractive conspecific individuals, via a “magnet effect.” Group attractiveness is thus potentially a public good that can be exploited by individuals, with selfish exploitation predicted to depend on genetic relatedness within the group. Petal colour is a potentially costly trait involved in floral signalling and advertising to pollinators. Here, we assessed whether petal colour was plastically sensitive to the relatedness of neighbours in the annual herb Moricandia moricandioides, which produces purple petals through anthocyanin pigment accumulation. We also tested whether petal colour intensity was related to nectar volume and sugar content in a context-dependent manner. Although both petal colour and petal anthocyanin concentration did not significantly vary with the neighbourhood configuration, plants growing with kin made a significantly higher investment in petal anthocyanin pigments as a result of the greater number and larger size of their flowers. Moreover the genetic relatedness of neighbours significantly modified the relationship between floral signalling and reward quantity: while focal plants growing with non-kin showed a positive relationship between petal colour and nectar production, plants growing with kin showed a positive relationship between number of flowers and nectar volume, and sugar content. The observed plastic response to group relatedness might have important effects on pollinator behaviour and visitation, with direct and indirect effects on plant reproductive success and mating patterns, at least in those plant species with patchy and genetically structured populations.

Authors thank Raquel Sánchez, Angel Caravante, Isabel Sánchez Almazo, Tatiana López Pérez, Samuel Cantarero, María José Jorquera and Germán Fernández for helping us during several phases of the study and Iván Rodríguez Arós for drawing the insect silhouettes. This research is supported by grants from the Spanish Ministry of Science, Innovation and Universities (CGL2015-71634-P, CGL2015-63827-P, CGL2017-86626-C2-1-P, CGL2017- 86626-C2-2-P, UNGR15-CE-3315, including EU FEDER funds), Junta de Andalucía (P18- FR-3641), Xunta de Galicia (CITACA), BBVA Foundation (PR17_ECO_0021), and a contract grant to C.A. from the former Spanish Ministry of Economy and Competitiveness (RYC-2012-12277). This is a contribution to the Research Unit Modeling Nature, funded by the Consejería de Economía, Conocimiento, Empresas y Universidad, and European Regional Development Fund (ERDF), reference SOMM17/6109/UGR., Phenotypic plasticity, the ability of a genotype of producing different phenotypes when exposed to different environments, may impact ecological interactions. We study here how within-individual plasticity in Moricandia arvensis flowers modifies its pollination niche. During spring, this plant produces large, cross-shaped, UV-reflecting lilac flowers attracting mostly long-tongued large bees. However, unlike most co-occurring species, M. arvensis keeps flowering during the hot, dry summer due to its plasticity in key vegetative traits. Changes in temperature and photoperiod in summer trigger changes in gene expression and the production of small, rounded, UV-absorbing white flowers that attract a different assemblage of generalist pollinators. This shift in pollination niche potentially allows successful reproduction in harsh conditions, facilitating M. arvensis to face anthropogenic perturbations and climate change. Floral phenotypes impact interactions between plants and pollinators. Here, the authors show that Moricandia arvensis displays discrete seasonal plasticity in floral phenotype, with large, lilac flowers attracting long-tongued bees in spring and small, rounded, white flowers attracting generalist pollinators in summer., Spanish Ministry of Science, Innovation and Universities (EU FEDER funds) CGL2015-71634-P CGL2015-63827-P CGL2017-86626-C2-1-P CGL2017-86626-C2-2-P UNGR15-CE-3315, Junta de Andalucia P18-FR-3641, Xunta de Galicia, BBVA Foundation PR17_ECO_0021, Spanish Ministry of Economy and Competitiveness RYC-2012-12277, Consejeria de Economia, Conocimiento, Empresas y Universidad SOMM17/6109/UGR, European Union (EU) SOMM17/6109/UGR

The study of intraspecific seed packaging (i.e. seed size/number strategy) variation across different populations may allow better understanding of the ecological forces that drive seed evolution in plants. Juniperus thurifera (Cupressaceae) provides a good model to study this due to the existence of two subspecies differentiated by phenotypic traits, such as seed size and cone seediness (number of seeds inside a cone), across its range. The aim of this study was to analyse seed packaging (seed mass and cone seediness) variation at different scales (subspecies, populations and individuals) and the relationship between cone and seed traits in European and African J. thurifera populations. After opening more than 5300 cones and measuring 3600 seeds, we found that seed packaging traits followed different patterns of variation. Large-scale effects (region and population) significantly contributed to cone seediness variance, while most of the seed mass variance occurred within individuals. Seed packaging differed between the two sides of the Mediterranean Sea, with African cones bearing fewer but larger seeds than the European ones. However, no differences in seed mass were found between populations when taking into account cone seediness. Larger cones contained more pulp and seeds and displayed a larger variation in individual seed mass. We validated previous reports on the intraspecific differences in J. thurifera seed packaging, although both subspecies followed the same seed size/number trade-off. The higher seediness and variation in seed mass found in larger cones reveals that the positive relationship between seed and cone sizes may not be straightforward.We hypothesise that the large variation of seed size found within cones and individuals in J. thurifera, but also in other fleshy-fruited species, could represent a bet-hedging strategy for dispersal.

The presence of visible annual rings in semi-arid tropical trees may allow the application of dendrochronological methods. However, variation in water availability may cause the formation of narrow, irregular ill- or non-defined annual rings hindering the correct dating of tree-ring series. We aimed to evaluate X-ray densitometry as a method to identify tree rings of two deciduous tree species from the Caatinga forest, a semi-arid region in the northeast of Brazil, and compare with two other methods commonly used in dendrochronology, the sliding-stage micrometer and image analysis. Xylem was observed macro- and microscopically and wood anatomical features were assessed in Aspidosperma pyrifolium and Poincianella pyramidalis trees. In both species, tree-ring boundaries were identified considering intra-annual density patterns and wood anatomical features. No significant differences in tree-ring widths were found among methods. X-ray densitometry measurements showed a positive correlation with the measurements obtained with image analysis and sliding-stage micrometer in A. pyrifolium and P. pyramidalis, revealing the high reliability of the methods used. However, inter-correlation of tree-ring width series showed differences in the accuracy of crossdating across measuring methods. The maximum, mean and minimum density values were species-dependent, with mean wood density of A. pyrifolium lower than P. pyramidalis. Our results highlight X-ray densitometry as an important and complementary tool to identify tree-rings boundaries in semi-arid tree species, especially in A. pyrifolium. Along with other measuring methods, it may provide higher accuracy in dendrochronological studies in semi-arid or subtropical environments.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality., Resilience to drought is crucial for tree survival under climate change. Here, DeSoto et al. show that trees that died during drought were less resilient to previous dry events compared to surviving conspecifics, but the resilience strategies differ between angiosperms and gymnosperms.

Exploring features of wood anatomy associated with fire scars found on fossil tree trunks is likely to increase our knowledge of the environmental and ecological processes that occurred in ancient forests and of the role of fire as an evolutionary force. In Petrified Forest National Park, Arizona, where Late Triassic fossil trees are exposed, we found 13 examples of fossil logs with external features resembling modern fire scars. One specimen with the unambiguous external features of a fire scar was collected for analysis of its fossilized wood. A light-colored band composed of compressed and distorted tracheids was associated with the scarring event. Cell lumen diameter and cell wall thickness in the pre-scarring fossilized wood show a response similar to that described in modern trees experiencing drought conditions. Tracheids in the post-scarring wood are initially smaller, and then become larger than average following a recovery period, as is often observed in modern conifers following fire. The responses in external morphology and wood anatomy to drought and fire were similar to those of some modern trees and support the view that some forests may have experienced conditions favoring the evolution of fire-adapted traits for more than 200 million years.

Fire scars are well known to fire ecologists and dendrochronologists worldwide, and are used in dating fires and reconstructing the fire histories of modern forests. Evidence of fires in ancient forests, such as fossil charcoal (fusain), is well known to paleontologists and has been reported in geologic formations dating back to the Late Devonian. We describe what we conclude is a fire scar on a fossil tree trunk from the Late Triassic Chinle Formation of southeastern Utah (~ 200–225 Ma). The external features of the prehistoric scar match those of modern fire scars better than those of scars created by other kinds of wounding events. The fossil specimen also exhibits a number of changes in wood anatomy similar to those reported in modern fire-scarred trees, including a band of very small tracheids that indicate growth suppression immediately associated with the scarring event; an area with a tangential row of probable traumatic resin ducts; and a significant increase in tracheid size following the scarring event that indicates a growth release. No fire scar resembling those in modern trees has previously been described in petrified wood as far as we can determine. The presence of a fire scar not only provides further evidence of ancient fires, but also shows that at least some individual trees survived them, indicating that fire could have been an ecological and evolutionary force in forests at least as early as the Late Triassic.

Because climate can affect xylem cell anatomy, series of intra-annual cell anatomical features have the potential to retrospectively supply seasonal climatic information. In this study, we explored the ability to extract information about water stress conditions from tracheid features of the Mediterranean conifer Juniperus thurifera L. Tracheidograms of four climatic years from two drought-sensitive sites in Spain were compared to evaluate whether it is possible to link intra-annual cell size patterns to seasonal climatic conditions. Results indicated site-specific anatomical adjustment such as smaller and thicker tracheids at the dryer site but also showed a strong climatic imprint on the intra-annual pattern of tracheid size. Site differences in cell size reflected expected structural adjustments against cavitation failures. Differences between intra-annual patterns, however, indicated a response to seasonal changes in water availability whereby cells formed under drought conditions were smaller and thicker, and vice versa. This relationship was more manifest and stable at the dryer site.

Plant–plant interactions are largely influenced by both environmental stress and ontogeny. Despite the effects of each of these factors on the overall outcome of these interactions has received considerable attention during the last years, the joint effects of both factors as drivers of such outcome are poorly understood. We used the combination of spatial pattern analysis, fruit production surveys, carbohydrate assays, sowing experiments and dendrochronological techniques to explore the interaction between Stipa tenacissima (nurse) and Lepidium subulatum (protegee) in two different slope aspects. This battery of techniques allows us to study the effects of the nurse plant during the whole life cycle of the protegee, and to assess the role of spatio-temporal variability in abiotic stress as a modulator of ontogenetic shifts in plant–plant interactions. Spatial pattern analyses suggested a net facilitative effect of S. tenacissima on L. subulatum . This effect was particularly important during the germination, shifting to competition (growth reduction) early after establishment. Competition was gradually reduced as the shrub aged, suggesting niche differentiation. The magnitude of competition was reduced under low rainfall levels in south-facing slopes, whereas this response was observed due to other abiotic factors in north-facing slopes. Our results highlight the crucial effect that positive interactions at early life-stages have to determine the long-term outcome of a given plant–plant interaction, and the existence of multiple shifts between facilitation and competition along different life-stages of the protegee. They also show how these ontogenetic shifts are modulated by abiotic factors, which differ among slope aspects. These findings may help to refine conceptual and theoretical models about shifts between facilitation and ontogeny under current climate change scenarios.

Tree features may modulate the sensitivity of radial growth to climate, leading to a nonuniform response. Age-related increases in climatic sensitivity have been observed repeatedly. Sex-related climatic sensitivity is also possible because of the long-term differential reproductive cost between the sexes. This study analysed the simultaneous effects of age and sex on the sensitivity of tree-ring growth to climate. Ring widths were measured from 50 female and 50 male Juniperus thurifera trees, 50-350 yr old, growing under a Mediterranean continental climate. Response functions were calculated based on tree-ring chronologies and monthly climatic records. Climatic sensitivity decreased with increasing age. Young trees (50-100 yr) were the most climatically sensitive to June-July precipitation, which affected growth positively. We found a significant interaction between age and sex in the climatic response of J. thurifera, with young females the most sensitive to summer water stress. Our results suggest that age-dependent climatic sensitivity can be determined by site-specific limiting environmental conditions and species-specific architectural and physiological adjustments during ontogeny. This study supports that the different ontogenetic stages of J. thurifera differ in their root structural traits and that sex-related sensitivity to summer drought may be attributable to less efficient water use by females.

Summary 1. In environmental sex determination (ESD) gender is decided after conception, depending on the environment, rather than being genetically fixed. ESD in plants has been mainly studied in angiosperms, where the dominant form of ESD is sexual lability. Surprisingly, ESD has rarely been studied in homosporous ferns, the only plants in which ESD is the rule, rather than the exception. 2. Here, we address the mechanism underlying ESD for the fern Woodwardia radicans by experimentally manipulating nutrient availability and gametophyte density. 3. Stress (limited nutrient supply, crowding) affected sexual expression in W. radicans gametophytes in a way compatible with ESD. Under good growth conditions (low density or high nutrient), gametophytes matured sexually at a relatively large size and turned into females and subsequently into bisexuals. Under harsh growth conditions, gametophytes matured sexually at a smaller size and turned into males. 4. Interestingly, gametophyte sexual expression was consistent with the size-advantage model, because the number of archegonia increased with gametophyte size, but not the number of antheridia. 5. The sex switch threshold size was variable and decreased with increasing stress, as predicted by age and size to maturity models. 6. Synthesis. Sexual expression in fern gametophytes can be fruitfully studied within the ESD theoretical framework. Stress induced male expression in gametophytes in a way compatible with ESD. In addition, size-related patterns of sexual expression were consistent with the size-advantage model, because female function benefited more from a larger size than did male function. Finally, the sex switch threshold size was variable and decreased with increasing stress, a result predicted by age and size to maturity models but seldom empirically tested before.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1–100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks., GGI by the Spanish Ministry of Economy and Competitiveness (Project AGL2014-61175-JIN)., AMH by the Spanish Ministry of Science and Innovation (Projects CGL2007-60120 and CSD2008-0040) and by the Spanish Ministry of Education via a FPU Scholarship., JJC, JCLC, and GSB by the Spanish Ministry of Economy (Projects CGL2015-69186-C2-1-R, CGL2013-48843-C2-2-R, and CGL2012-32965) and the EU (Project FEDER 0087 TRANSHABITAT).

Differences in reproductive costs between male and female plants have been shown to foster sex-related variability in growth and C-storage patterns. The extent to which differential secondary growth in dioecious trees is associated with changes in stem carbohydrate storage patterns, however, has not been fully assessed. We explored the long-term radial growth and the seasonal variation of non-structural carbohydrate (NSC) content in sapwood of 40 male and 40 female Juniperus thurifera trees at two sites. NSC content was analyzed bimonthly for one year, and tree-ring width was measured for the 1931–2010 period. Sex-related differences in secondary growth and carbohydrate storage differed between sites. Under less restrictive environmental conditions females grew more and stored more non-soluble sugars than males. Our results reinforce that sex-related differences in growth and resource storage may be a consequence of local adaptation to environmental conditions. Seasonal variation in soluble sugars concentration was opposite to cambial activity, with minima seen during periods of maximal secondary growth, and did not differ between the sexes or sites. Trees with higher stem NSC levels at critical periods showed higher radial growth, suggesting a common mechanism irrespective of site or sex. Sex-related patterns of secondary growth were linked to differences in non-soluble sugars content indicating sex-specific strategies of long-term performance.

This paper was commissioned by the members of the Ecosphere Editorial Board to commemorate the ESA Centennial celebration.

An increased understanding of intraspecific seed packaging (i.e. seed size/number strategy) variation across different environments may improve current knowledge of the ecological forces that drive seed evolution in plants. In particular, pre-dispersal seed predation may influence seed packaging strategies, triggering a reduction of the resources allocated to undamaged seeds within the preyed fruits. Assessing plant reactions to pre-dispersal seed predation is crucial to a better understanding of predation effects, but the response of plants to arthropod attacks remains unexplored. We have assessed the effect of cone predation on the size and viability of undamaged seeds in populations of Juniperus thurifera with contrasting seed packaging strategies, namely, North African populations with single-large-seeded cones and South European populations with multi-small-seeded cones. Our results show that the incidence of predation was lower on the single-large-seeded African cones than on the multi-small-seeded European ones. Seeds from non-preyed cones were also larger and had a higher germination success than uneaten seeds from preyed cones, but only in populations with multi-seeded cones and in cones attacked by Trisetacus sp., suggesting a differential plastic response to predation. It is possible that pre-dispersal seed predation has been a strong selective pressure in European populations with high cone predation rates, being a process which maintains multi-small-seeded cones and empty seeds as a strategy to save some seeds from predation. Conversely, pre-dispersal predation might not have a strong effect in the African populations with single-large-seeded cones characterized by seed germination and filling rates higher than those in the European populations. Our results indicate that differences in pre-dispersal seed predators and predation levels may affect both selection on and intraspecific variation in seed packaging.

Geographically structured tree-ring networks are needed to fully understand the spatiotemporal variability in climatic sensitiveness of trees and to study their future responses to global warming. We aim to identify the spatially constrained structure of radial-growth patterns of the Spanish juniper (Juniperus thurifera L.) and to assess whether their climate–growth responses were unstable during the late twentieth century. Tree–ring width chronologies were built for 13 J. thurifera stands in Spain using dendrochronological methods and related to monthly climatic data. Sites were grouped according to their growth patterns using hierarchical cluster analysis. The relationships among geographical, climatic and stand features and their influence on radial growth were evaluated using redundancy analysis. The climate–growth relationships and their temporal stability were assessed using Pearson’s and moving bootstrapped correlations, respectively. Stands formed three geographical groups according to their highfrequency growth variation: North West and Centre, North East and South East. We found that J. thurifera radialgrowth patterns depended on geographical and climatic factors, but not on the stand structure, and responded to a northwest–southeast gradient of decreasing rainfall and influence of Atlantic Westerlies and Mediterranean cyclonic activity. The positive response to June precipitation was unstable during the late twentieth century and started earlier in populations from western mesic sites than in eastern xeric sites. This pattern may be related to either decreasing water availability in western than in eastern sites or the resilience of J. thurifera growth from xeric sites in response to the increasing summer aridity., Junta de Castilla y León supported this research with the projects VA069A07, VA006A10-2 and MEDIATIC (PTDC/AAC-CLI/ 103361/2008).

• Juniper woodlands have a great conservational and productive importance in Spain. However, basic traits of their structural variation and growth dynamics are unknown. • To characterize the structural variation and assess both height and radial growth patterns, we sampled and measured stem cross-sections every 0.5 m in height from 107 Spanish juniper (Juniperus thurifera L.) trees, which were selected based on a 350 m regular grid over a whole woodland of 3 300 ha in area. We used dendroecological techniques and spatial analysis to study tree structure and growth. • Structural traits such as bole diameter and height were mainly determined by tree life-related parameters such as tree age and growth rate variation, while abiotic factors such as altitude played a minor role in determining structural variation. Over the last 300 years, juniper establishment has been continuous in time but discontinuous in space. Large-scale spatial heterogeneity of tree establishment, and the presence of an early growth suppression and abrupt growth changes in junipers > 100 years old are consequences of an intense management that almost ceased in the late 19th century. • Abrupt growth releases and suppressions were synchronic among different age classes, and coincided with documentary records on livestock decrease and key historical changes in land use. Growth patterns suggest that juniper is a brow sing-tolerant species that is able to survive large periods of intense browsing pressure.

DeSoto, L., Torices, R., Broennimann, O., Guisan, A., Rodriguez-Echeverria, S. 2016. Is there a bias in participation and visibility against women in ecology? A comparison between Iberian and Swiss conferences. Ecosistemas 25(3):105-111. Doi.: 10.7818/ECOS.2016.25-3.12 Women are still under-represented in science and technology because there is a bias in the different evaluation processes from the admission of oral communications at conferences to the access to more qualified positions. In this study, we compared the participation and visibility of female ecologists in the IV Iberian Ecological Conference (CIE4) with the Annual Swiss Conference on Ecology, Evolution, Systematics, Biogeography and Conservation (Biology16) where a method of blind evaluation was applied. In both conferences, and after active selection by conference committees, male delegates presented proportionately more regular oral contributions than female ones, although these differences were marginally significant. In the Biology16, this bias was due to lower female applicants for oral contributions, and in the CIE4, to a lower selection rate of female oral contributions. In the CIE4, the higher male presence in oral contributions was consistent with male greater visibility, whereas female visibility was higher in the Biology16. This different visibility was caused by contrasting selection for invited speakers; being male researches preferentially invited at the CIE4, but female ones at the Biology16. Implementing a blind review system of contributions and active policies to promote the participation of women as invited speakers may reduce the differences in visibility and could contribute in the medium term to eliminate the bias against women in the selection of oral contributions at the Iberian conferences.

A recommendation of: Cathleen Petit-Cailleux, Hendrik Davi, Francois Lefevre, Christophe Hurson, Joseph Garrigue, Jean-Andre Magdalou, Elodie Magnanou and Sylvie Oddou-Muratorio Combining statistical and mechanistic models to identify the drivers of mortality within a rear-edge beech population 10.1101/645747