Long-term rain exclusion in a Mediterranean forest: response of physiological and physico-chemical traits of Quercus pubescens across seasons.

With climate change, an aggravation in summer drought is expected in the Mediterranean region. To assess the impact of such a future scenario, we compared the response of Quercus pubescens, a drought-resistant deciduous oak species, to long-term amplified drought (AD) (partial rain exclusion in natura for 10 years) and natural drought (ND). We studied leaf physiological and physico-chemical trait responses to ND and AD over the seasonal cycle, with a focus on chemical traits including major groups of central (photosynthetic pigments and plastoquinones) and specialized (tocochromanols, phenolic compounds, and cuticular waxes) metabolites. Seasonality was the main driver of all leaf traits, including cuticular triterpenoids, which were highly concentrated in summer, suggesting their importance to cope with drought and thermal stress periods. Under AD, trees not only reduced CO2 assimilation (-42%) in summer and leaf concentrations of some phenolic compounds and photosynthetic pigments (carotenoids from the xanthophyll cycle) but also enhanced the levels of other photosynthetic pigments (chlorophylls, lutein, and neoxanthin) and plastochromanol-8, an antioxidant located in chloroplasts. Overall, the metabolomic adjustments across seasons and drought conditions reinforce the idea that Q. pubescens is highly resistant to drought although significant losses of antioxidant defenses and photoprotection were identified under AD.

Genomic divergence and demographic history of Quercus aliena populations.

BACKGROUND: Quercus aliena is a major montane tree species of subtropical and temperate forests in China, with important ecological and economic value. In order to reveal the species' population dynamics, genetic diversity, genetic structure, and association with mountain habitats during the evolutionary process, we re-sequenced the genomes of 72 Q. aliena individuals. RESULTS: The whole chloroplast and nuclear genomes were used for this study. Phylogenetic analysis using the chloroplast genome dataset supported four clades of Q. aliena, while the nuclear dataset supported three major clades. Sex-biased dispersal had a critical role in causing discordance between the chloroplast and nuclear genomes. Population structure analysis showed two groups in Q. aliena. The effective population size sharply declined 1 Mya, coinciding with the Poyang Glaciation in Eastern China. Using genotype-climate association analyses, we found a positive correlation between allele frequency variation in SNPs and temperature, suggesting the species has the capacity to adapt to changing temperatures. CONCLUSION: Overall, this study illustrates the genetic divergence, genomic variation, and evolutionary processes behind the demographic history of Q. aliena.

Genomic insights into local adaptation and vulnerability of Quercus longinux to climate change.

BACKGROUND: Climate change is expected to alter the factors that drive changes in adaptive variation. This is especially true for species with long life spans and limited dispersal capabilities. Rapid climate changes may disrupt the migration of beneficial genetic variations, making it challenging for them to keep up with changing environments. Understanding adaptive genetic variations in tree species is crucial for conservation and effective forest management. Our study used landscape genomic analyses and phenotypic traits from a thorough sampling across the entire range of Quercus longinux, an oak species native to Taiwan, to investigate the signals of adaptation within this species. RESULTS: Using ecological data, phenotypic traits, and 1,933 single-nucleotide polymorphisms (SNPs) from 205 individuals, we classified three genetic groups, which were also phenotypically and ecologically divergent. Thirty-five genes related to drought and freeze resistance displayed signatures of natural selection. The adaptive variation was driven by diverse environmental pressures such as low spring precipitation, low annual temperature, and soil grid sizes. Using linear-regression-based methods, we identified isolation by environment (IBE) as the optimal model for adaptive SNPs. Redundancy analysis (RDA) further revealed a substantial joint influence of demography, geology, and environments, suggesting a covariation between environmental gradients and colonization history. Lastly, we utilized adaptive signals to estimate the genetic offset for each individual under diverse climate change scenarios. The required genetic changes and migration distance are larger in severe climates. Our prediction also reveals potential threats to edge populations in northern and southeastern Taiwan due to escalating temperatures and precipitation reallocation. CONCLUSIONS: We demonstrate the intricate influence of ecological heterogeneity on genetic and phenotypic adaptation of an oak species. The adaptation is also driven by some rarely studied environmental factors, including wind speed and soil features. Furthermore, the genetic offset analysis predicted that the edge populations of Q. longinux in lower elevations might face higher risks of local extinctions under climate change.

An improved method to study Phytophthora cinnamomi Rands zoospores interactions with host.

Phytophthora cinnamomi Rands is a highly prevalent phytopathogen worldwide, ranking among the top ten in terms of distribution. It inflicts crown rot, canker, and root rot on numerous plant species, significantly impacting the biodiversity of both flora and fauna within affected environments. With a host range spanning over 5,000 species, including important plants like Quercus suber, Quercus ilex, Castanea sativa, and commercially significant crops such as avocado (Persea americana), maize (Zea mays), and tomato (Solanum lycopersicum), Phytophthora cinnamomi poses a substantial threat to agriculture and ecosystems. The efficient dissemination of the oomycete relies on its short-lived asexually motile zoospores, which depend on water currents to infect host roots. However, managing these zoospores in the laboratory has long been challenging due to the complexity of the life cycle. Current protocols involve intricate procedures, including alternating cycles of growth, drought, and flooding. Unfortunately, these artificial conditions often result in a rapid decline in virulence, necessitating additional steps to maintain infectivity during cultivation. In our research, we sought to address this challenge by investigating zoospore survival under various conditions. Our goal was to develop a stable stock of zoospores that is both easily deployable and highly infective. Through direct freezing in liquid nitrogen, we have successfully preserved their virulence. This breakthrough eliminates the need for repeated culture transfers, simplifying the process of plant inoculation. Moreover, it enables more comprehensive studies of Phytophthora cinnamomi and its interactions with host plants.

Leaf ecological stoichiometry and anatomical structural adaptation mechanisms of Quercus sect. Heterobalanus in southeastern Qinghai-Tibet Plateau.

BACKGROUND: With the dramatic uplift of the Qinghai-Tibet Plateau (QTP) and the increase in altitude in the Pliocene, the environment became dry and cold, thermophilous plants that originally inhabited ancient subtropical forest essentially disappeared. However, Quercus sect. Heterobalanus (QSH) have gradually become dominant or constructive species distributed on harsh sites in the Hengduan Mountains range in southeastern QTP, Southwest China. Ecological stoichiometry reveals the survival strategies plants adopt to adapt to changing environment by quantifying the proportions and relationships of elements in plants. Simultaneously, as the most sensitive organs of plants to their environment, the structure of leaves reflects of the long-term adaptability of plants to their surrounding environments. Therefore, ecological adaptation mechanisms related to ecological stoichiometry and leaf anatomical structure of QSH were explored. In this study, stoichiometric characteristics were determined by measuring leaf carbon (C), nitrogen (N), and phosphorus (P) contents, and morphological adaptations were determined by examining leaf anatomical traits with microscopy. RESULTS: Different QSH life forms and species had different nutrient allocation strategies. Leaves of QSH plants had higher C and P and lower N contents and higher N and lower P utilization efficiencies. According to an N: P ratio threshold, the growth of QSH species was limited by N, except that of Q. aquifolioides and Q. longispica, which was limited by both N and P. Although stoichiometric homeostasis of C, N, and P and C: N, C: P, and N: P ratios differed slightly across life forms and species, the overall degree of homeostasis was strong, with strictly homeostatic, homeostatic, and weakly homeostatic regulation. In addition, QSH leaves had compound epidermis, thick cuticle, developed palisade tissue and spongy tissue. However, leaves were relatively thin overall, possibly due to leaf leathering and lignification, which is strategy to resist stress from UV radiation, drought, and frost. Furthermore, contents of C, N, and P and stoichiometric ratios were significantly correlated with leaf anatomical traits. CONCLUSIONS: QSH adapt to the plateau environment by adjusting the content and utilization efficiencies of C, N, and P elements. Strong stoichiometric homeostasis of QSH was likely a strategy to mitigate nutrient limitation. The unique leaf structure of the compound epidermis, thick cuticle, well-developed palisade tissue and spongy tissue is another adaptive mechanism for QSH to survive in the plateau environment. The anatomical adaptations and nutrient utilization strategies of QSH may have coevolved during long-term succession over millions of years.

Hydrogen isotope fractionation in plants with C3, C4, and CAM CO2 fixation.

Measurements of stable isotope ratios in organic compounds are widely used tools for plant ecophysiological studies. However, the complexity of the processes involved in shaping hydrogen isotope values (δ2H) in plant carbohydrates has limited its broader application. To investigate the underlying biochemical processes responsible for 2H fractionation among water, sugars, and cellulose in leaves, we studied the three main CO2 fixation pathways (C3, C4, and CAM) and their response to changes in temperature and vapor pressure deficit (VPD). We show significant differences in autotrophic 2H fractionation (εA) from water to sugar among the pathways and their response to changes in air temperature and VPD. The strong 2H depleting εA in C3 plants is likely driven by the photosynthetic H+ production within the thylakoids, a reaction that is spatially separated in C4 and strongly reduced in CAM plants, leading to the absence of 2H depletion in the latter two types. By contrast, we found that the heterotrophic 2H-fractionation (εH) from sugar to cellulose was very similar among the three pathways and is likely driven by the plant's metabolism, rather than by isotopic exchange with leaf water. Our study offers new insights into the biochemical drivers of 2H fractionation in plant carbohydrates.

Ancient tree genomes for old questions.

Most foundational work on the evolution and migration of plant species relies on genomic data from contemporary samples. Ancient plant samples can give us access to allele sequences and distributions on the landscape dating back to the mid Holocene or earlier (Gugerli et al., 2005). Nuclear DNA from ancient wood, however, has been mostly inaccessible until now. In a From the Cover article in this issue of Molecular Ecology, Wagner et al. (2023) present the first resequenced nuclear genomes from ancient oak wood, including two samples dated to the 15th century and one that dates to more than 3500 years ago. These ancient tree genomes open the possibility for investigating species adaptation, migration, divergence, and hybridisation in the deep past. They pave the way for what we hope will be a new era in the use of paleogenomics to study Holocene tree histories.

Genetic differentiation across a steep and narrow environmental gradient: Quantitative genetic and genomic insights into Lake Superior populations of Quercus rubra.

Adaptive differentiation of traits and underlying loci can occur at a small geographical scale if natural selection is stronger than countervailing gene flow and drift. We investigated this hypothesis using coupled quantitative genetic and genomic approaches for a wind-pollinated tree species, Quercus rubra, along the steep, narrow gradient of the Lake Superior coast that encompasses four USDA Hardiness Zones within 100 km. For the quantitative genetic component of this study, we examined phenotypic differentiation among eight populations in a common garden, measuring seed mass, germination, height, stem diameter, leaf number, specific leaf area and survival. For the genomic component, we quantified genetic differentiation for 26 populations from the same region using RAD-seq. Because hybridisation with Quercus ellipsoidalis occurs in other parts of the species' range, we included two populations of this congener for comparison. In the common garden study, we found a strong signal of population differentiation that was significantly associated with at least one climate factor for nine of 10 measured traits. In contrast, we found no evidence of genomic differentiation among populations based on FST or any other measures. However, both distance-based and genotype-environment association analyses identified loci showing the signature of selection, with one locus in common across five analyses. This locus was associated with the minimum temperature of the coldest month, a factor that defines the climate zones and was also significant in the common garden analyses. In addition, we documented introgression from Q. ellipsoidalis into Q. rubra, with rates of introgression correlated with the climate gradient. In sum, this study reveals signatures of selection at the quantitative trait and genomic level consistent with climate adaptation, a pattern that is more often documented at a much broader geographical scale, especially in long-lived wind-pollinated species.

Tracking population structure and phenology through time using ancient genomes from waterlogged white oak wood.

Whole genome characterizations of crop plants based on ancient DNA have provided unique keys for a better understanding of the evolutionary origins of modern cultivars, the pace and mode of selection underlying their adaptation to new environments and the production of phenotypes of interest. Although forests are among the most biologically rich ecosystems on earth and represent a fundamental resource for human societies, no ancient genome sequences have been generated for trees. This contrasts with the generation of multiple ancient reference genomes for important crops. Here, we sequenced the first ancient tree genomes using two white oak wood remains from Germany dating to the Last Little Ice Age (15th century CE, 7.3× and 4.0×) and one from France dating to the Bronze Age (1700 BCE, 3.4×). We assessed the underlying species and identified one medieval remains as a hybrid between two common oak species (Quercus robur and Q. petraea) and the other two remains as Q. robur. We found that diversity at the global genome level had not changed over time. However, exploratory analyses suggested that a reduction of diversity took place at different time periods. Finally, we determined the timing of leaf unfolding for ancient trees for the first time. The study extends the application of ancient wood beyond the classical proxies of dendroclimatology, dendrochronology, dendroarchaeology and dendroecology, thereby enhancing resolution of inferences on the responses of forest ecosystems to past environmental changes, epidemics and silvicultural practices.

Restricted internal diffusion weakens transpiration-photosynthesis coupling during heatwaves: Evidence from leaf carbonyl sulphide exchange.

Increasingly frequent and intense heatwaves threaten ecosystem health in a warming climate. However, plant responses to heatwaves are poorly understood. A key uncertainty concerns the intensification of transpiration when heatwaves suppress photosynthesis, known as transpiration-photosynthesis decoupling. Field observations of such decoupling are scarce, and the underlying physiological mechanisms remain elusive. Here, we use carbonyl sulphide (COS) as a leaf gas exchange tracer to examine potential mechanisms leading to transpiration-photosynthesis decoupling on a coast live oak in a southern California woodland in spring 2013. We found that heatwaves suppressed both photosynthesis and leaf COS uptake but increased transpiration or sustained it at non-heatwave levels throughout the day. Despite statistically significant decoupling between transpiration and photosynthesis, stomatal sensitivity to environmental factors did not change during heatwaves. Instead, midday photosynthesis during heatwaves was restricted by internal diffusion, as indicated by the lower internal conductance to COS. Thus, increased evaporative demand and nonstomatal limitation to photosynthesis act jointly to decouple transpiration from photosynthesis without altering stomatal sensitivity. Decoupling offered limited potential cooling benefits, questioning its effectiveness for leaf thermoregulation in xeric ecosystems. We suggest that adding COS to leaf and ecosystem flux measurements helps elucidate diverse physiological mechanisms underlying transpiration-photosynthesis decoupling.

Major tree species of Central European forests differ in their proportion of positive, negative, and nonstationary growth trends.

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree-ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990-2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation-based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected.

Responses of oak seedlings to increased herbivory and drought: a possible trade-off?

BACKGROUND AND AIMS: Anthropogenic disturbances are causing a co-occurring increase in biotic (ungulate herbivory) and abiotic (drought) stressors, threatening plant reproduction in oak-dominated ecosystems. However, we wonder whether herbivory could compensate for the adverse impact of drought by reducing evapotranspiration. Thus, we investigate the isolated and joint effects of herbivory and drought on oak seedlings of two contrasting Mediterranean species that differ in leaf habit and drought resistance. METHODS: California oak seedlings from the evergreen, and more drought-resistant, Quercus agrifolia and the deciduous Q. lobata (n=387) were assigned to a fully crossed factorial design with herbivory and drought as stress factors. Seedlings were assigned in a greenhouse to 3-4 clipping levels simulating herbivory and 3-4 watering levels, depending on the species. We measured survival, growth, and leaf attributes (chlorophyll, secondary metabolites, leaf area and weight) once a month (May-Sep) and harvested above- and below-ground biomass at the end of the growing season. KEY RESULTS: For both oak species, simulated herbivory enhanced seedling survival during severe drought or delayed its adverse effects, probably due to reduced transpiration resulting from herbivory-induced leaf area reduction and compensatory root growth. Seedlings from the deciduous, and less drought-resistant species, benefitted from herbivory at lower levels of water stress, suggesting different response across species. We also found complex interactions between herbivory and drought on their impact on leaf attributes. In contrast to chlorophyll content which was not affected by herbivory, anthocyanins increased with herbivory - although water stress reduced differences in anthocyanins due to herbivory. CONCLUSIONS: Herbivory seems to facilitate Mediterranean oak seedlings to withstand summer drought, potentially alleviating a key bottleneck in the oak recruitment process. Our study highlights the need to consider ontogenetic stages and species-specific traits in understanding complex relationships between herbivory and drought stressors for the persistence and restoration of multi-species oak savannas.

Divergence and reticulation in the Mexican white oaks: ecological and phylogenomic evidence on species limits and phylogenetic networks in the Quercus laeta complex (Fagaceae).

BACKGROUND AND AIMS: Introgressive hybridization poses a challenge to taxonomic and phylogenetic understanding of taxa, particularly when there are high numbers of co-occurring, intercrossable species. The genus Quercus exemplifies this situation. Oaks are highly diverse in sympatry and cross freely, creating syngameons of interfertile species. Although a well-resolved, dated phylogeny is available for the American oak clade, evolutionary relationships within many of the more recently derived clades remain to be defined, particularly for the young and exceptionally diverse Mexican white oak clade. Here, we adopted an approach bridging micro- and macroevolutionary scales to resolve evolutionary relationships in a rapidly diversifying clade endemic to Mexico. METHODS: Ecological data and sequences of 155 low-copy nuclear genes were used to identify distinct lineages within the Quercus laeta complex. Concatenated and coalescent approaches were used to assess the phylogenetic placement of these lineages relative to the Mexican white oak clade. Phylogenetic network methods were applied to evaluate the timing and genomic significance of recent or historical introgression among lineages. KEY RESULTS: The Q. laeta complex comprises six well-supported lineages, each restricted geographically and with mostly divergent climatic niches. Species trees corroborated that the different lineages are more closely related to other species of Mexican white oaks than to each other, suggesting that this complex is polyphyletic. Phylogenetic networks estimated events of ancient introgression that involved the ancestors of three present-day Q. laeta lineages. CONCLUSIONS: The Q. laeta complex is a morphologically and ecologically related group of species rather than a clade. Currently, oak phylogenetics is at a turning point, at which it is necessary to integrate phylogenetics and ecology in broad regional samples to figure out species boundaries. Our study illuminates one of the more complicated of the Mexican white oak groups and lays groundwork for further taxonomic study.

Early overyielding in a mixed deciduous forest is driven by both above- and below ground species-specific acclimation.

BACKGROUND AND AIMS: Mixed forest plantations are increasingly recognised for their role in mitigating the impacts of climate change and enhancing ecosystem resilience. Yet, there remains a significant gap in understanding the early-stage dynamics of species trait diversity and interspecies interactions, particularly in pure deciduous mixtures. This study aims to explore the timing and mechanisms by which trait diversity of deciduous species and competitive interactions influence yield, carbon allocation, and space occupation in mixed forests, both above- and belowground. METHODS: A forest inventory was conducted in planted monocultures, 2-species, and 4-species mixtures of European Acer, Tilia, Carpinus, and Quercus, representing a spectrum from acquisitive to conservative tree species. Competition effects were assessed with linear mixed-effects models at the level of biomass and space acquisition, including leaf, canopy, stem, and fine root traits. KEY RESULTS: Early aboveground growth effects were observed six years post-planting, with significant biomass accumulation after eight years, strongly influenced by species composition. Mixtures, especially with acquisitive species, exhibited aboveground overyielding, 1.5- to 1.9-times higher than monocultures. Fine roots showed substantial overyielding in high diversity stands. Biomass allocation was species-specific and varied markedly by tree size, the level of diversity, and between acquisitive Acer and the more conservative species. No root segregation was found. CONCLUSIONS: Our findings underscore the critical role of species trait diversity in enhancing productivity in mixed deciduous forest plantations. Allometric changes highlight the need to differentiate between (active) acclimations and (passive) tree size-related changes, but illustrate major consequences of competitive interactions for the functional relation between leaves, stem, and roots. This study points towards the significant contributions of both above- and belowground components to overall productivity of planted mixed-species forests.

Offspring may succeed well next to their relatives, but it needs particular traits.

BACKGROUND AND AIMS: There is ongoing debate about whether offspring perform best next to phylogenetically distantly related adult neighbours (due to the scarcity of enemies and competitors) or next to closely related adults (due to the abundance of mutualists). Here we hypothesise that relatedness of adult neighbours affects which traits confer performance rather than performance itself. METHODS: We studied seed removal, seed germination and sapling growth in Sessile Oaks (Quercus petraea and hybrids), and how they depend on size, shape and other traits, under both closely and distantly related canopies, manipulating offspring-density, presence of insects, and fungi, and spatial proximity to oaks. KEY RESULTS: We found that phylogenetic distance of adult neighbours affects only little performance of offspring but strongly which traits confer performance to offspring, in particular the size and shape of seeds and saplings. Differences in trait-performance relationships mostly disappear once insects or conspecific competitors are excluded (albeit exclusion of fungi reinforced these differences). Effects of phylogenetic distance of neighbours were not replaceable by the percentage of the gymnosperms among neighbours, nor the environmental conditions considered. CONCLUSIONS: We suggest that by responding to a biotic micro-mosaic of selection pressures, Sessile Oak flexibly succeeds in diverse neighbourhoods. Sessile Oak might maintain the potential for both, convergence with and divergence from phylogenetically distantly related species, thereby reinforcing or eroding phylogenetic signal of niches.

Introgression and incomplete lineage sorting blurred phylogenetic relationships across the genomes of sclerophyllous oaks from southwest China.

Resolving evolutionary relationships among closely related species with interspecific gene flow is challenging. Genome-scale data provide opportunities to clarify complex evolutionary relationships in closely related species and to observe variations in species relationships across the genomes of such species. The Himalayan-Hengduan subalpine oaks have a nearly completely sympatric distribution in southwest China and probably constitute a syngameon. In this study, we mapped resequencing data from different species in this group to the Quercus aquifolioides reference genome to obtain a high-quality filtered single nucleotide polymorphism (SNP) dataset. We also assembled their plastomes. We reconstructed their phylogenetic relationships, explored the level and pattern of introgression among these species and investigated gene tree variation in the genomes of these species using sliding windows. The same or closely related plastomes were found to be shared extensively among different species within a specific geographical area. Phylogenomic analyses of genome-wide SNP data found that most oaks in the Himalayan-Hengduan subalpine clade showed genetic coherence, but several species were found to be connected by introgression. The gene trees obtained using sliding windows showed that the phylogenetic relationships in the genomes of oaks are highly heterogeneous and therefore highly obscured. Our study found that all the oaks of the Himalayan-Hengduan subalpine clade from southwest China form a syngameon. The obscured phylogenetic relationships observed empirically across the genome are best explained by interspecific gene flow in conjunction with incomplete lineage sorting.

Ungulates mitigate the effects of drought and shrub encroachment on the fire hazard of Mediterranean oak woodlands.

Climate change is increasing the frequency of droughts and the risk of severe wildfires, which can interact with shrub encroachment and browsing by wild ungulates. Wild ungulate populations are expanding due, among other factors, to favorable habitat changes resulting from land abandonment or land-use changes. Understanding how ungulate browsing interacts with drought to affect woody plant mortality, plant flammability, and fire hazard is especially relevant in the context of climate change and increasing frequency of wildfires. The aim of this study is to explore the combined effects of cumulative drought, shrub encroachment, and ungulate browsing on the fire hazard of Mediterranean oak woodlands in Portugal. In a long-term (18 years) ungulate fencing exclusion experiment that simulated land abandonment and management neglect, we investigated the population dynamics of the native shrub Cistus ladanifer, which naturally dominates the understory of woodlands and is browsed by ungulates, comparing areas with (no fencing) and without (fencing) wild ungulate browsing. We also modeled fire behavior in browsed and unbrowsed plots considering drought and nondrought scenarios. Specifically, we estimated C. ladanifer population density, biomass, and fuel load characteristics, which were used to model fire behavior in drought and nondrought scenarios. Overall, drought increased the proportion of dead C. ladanifer shrub individuals, which was higher in the browsed plots. Drought decreased the ratio of live to dead shrub plant material, increased total fuel loading, shrub stand flammability, and the modeled fire parameters, that is, rate of surface fire spread, fireline intensity, and flame length. However, total fuel load and fire hazard were lower in browsed than unbrowsed plots, both in drought and nondrought scenarios. Browsing also decreased the population density of living shrubs, halting shrub encroachment. Our study provides long-term experimental evidence showing the role of wild ungulates in mitigating drought effects on fire hazard in shrub-encroached Mediterranean oak woodlands. Our results also emphasize that the long-term effects of land abandonment can interact with climate change drivers, affecting wildfire hazard. This is particularly relevant given the increasing incidence of land abandonment.

Sustaining eastern oak forests: Synergistic effects of fire and topography on vegetation and fuels.

Across much of the eastern United States, oak forests are undergoing mesophication as shade-tolerant competitors become more abundant and suppress oak regeneration. Given the historical role of anthropogenic surface fires in promoting oak dominance, prescribed fire has become important in efforts to reverse mesophication and sustain oaks. In 2000 we established the Ohio Hills Fire and Fire Surrogate (FFS) study to examine whether repeated prescribed fire (Fire), mechanical partial harvest (Mech), and their combined application (Mech + Fire) reduced the dominance of subcanopy mesophytic competitors, increased the abundance of large oak-hickory advance regeneration, created a more diverse and productive ground-layer flora, and produced fuel beds more conducive to prescribed fire, reducing the risk of high-severity wildfire. Here we report on the ~20-year effects of treatments on vegetation and fuels and examine the support for interactive effects across a topographic-moisture and energy gradient. In general, we found that Fire and Mech + Fire treatments tended to reverse mesophication while the Mech-only treatment did not. The moderate and occasionally high-intensity fires resulted in effects that were ultimately very similar between the two fire treatments but were modulated by topography with increasing fire severity on drier sites. In particular, we found support for an interaction effect between treatment and topography on forest structure and tree regeneration responses. Fire generally reduced mesophytic tree density in the midstory and sapling strata across all site conditions, while leading to substantial gains in the abundance of large oak-hickory advance regeneration on dry and intermediate landscape positions. Fire also promoted ground-layer diversity and created compositionally distinct communities across all site conditions, primarily through the increased richness of native perennial herbs. However, the fire had limited effects on fine surface fuel loading and increased the loading of large woody fuels, potentially increasing the risk of high-severity wildfire during drought conditions. We conclude that two decades of repeated fires, with and without mechanical density reduction, significantly shifted the trajectory of mesophication across most of the landscape, particularly on dry and intermediate sites, highlighting the capacity of a periodic fire regime to sustain eastern oak forests and promote plant diversity but modulated by topography.

Urbanization exacerbates climate sensitivity of eastern United States broadleaf trees.

Tree growth is a key mechanism driving carbon sequestration in forest ecosystems. Environmental conditions are important regulators of tree growth that can vary considerably between nearby urban and rural forests. For example, trees growing in cities often experience hotter and drier conditions than their rural counterparts while also being exposed to higher levels of light, pollution, and nutrient inputs. However, the extent to which these intrinsic differences in the growing conditions of trees in urban versus rural forests influence tree growth response to climate is not well known. In this study, we tested for differences in the climate sensitivity of tree growth between urban and rural forests along a latitudinal transect in the eastern United States that included Boston, Massachusetts, New York City, New York, and Baltimore, Maryland. Using dendrochronology analyses of tree cores from 55 white oak trees (Quercus alba), 55 red maple trees (Acer rubrum), and 41 red oak trees (Quercus rubra) we investigated the impacts of heat stress and water stress on the radial growth of individual trees. Across our three-city study, we found that tree growth was more closely correlated with climate stress in the cooler climate cities of Boston and New York than in Baltimore. Furthermore, heat stress was a significant hindrance to tree growth in higher latitudes while the impacts of water stress appeared to be more evenly distributed across latitudes. We also found that the growth of oak trees, but not red maple trees, in the urban sites of Boston and New York City was more adversely impacted by heat stress than their rural counterparts, but we did not see these urban-rural differences in Maryland. Trees provide a wide range of important ecosystem services and increasing tree canopy cover was typically an important component of urban sustainability strategies. In light of our findings that urbanization can influence how tree growth responds to a warming climate, we suggest that municipalities consider these interactions when developing their tree-planting palettes and when estimating the capacity of urban forests to contribute to broader sustainability goals in the future.

The effect of local habitat and spatial connectivity on urban seed predation.

PREMISE: During the last centuries, the area covered by urban landscapes is increasing all over the world. Urbanization can change local habitats and decrease connectivity among these habitats, with important consequences for species interactions. While several studies have found a major imprint of urbanization on plant-insect interactions, the effects of urbanization on seed predation remain largely unexplored. METHODS: We investigated the relative impact of sunlight exposure, leaf litter, and spatial connectivity on predation by moth and weevil larvae on acorns of the pedunculate oak across an urban landscape during 2018 and 2020. We also examined whether infestations by moths and weevils were independent of each other. RESULTS: While seed predation varied strongly among trees, seed predation was not related to differences in sunlight exposure, leaf litter, or spatial connectivity. Seed predation by moths and weevils was negatively correlated at the level of individual acorns in 2018, but positively correlated at the acorn and the tree level in 2020. CONCLUSIONS: Our study sets the baseline expectation that urban seed predators are unaffected by differences in sunlight exposure, leaf litter, and spatial connectivity. Overall, our findings suggest that the impact of local and spatial factors on insects within an urban context may depend on the species guild. Understanding the impact of local and spatial factors on biodiversity, food web structure, and ecosystem functioning can provide valuable insights for urban planning and management strategies aimed at promoting urban insect diversity.